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Full text of "Bulletin of the Natural History Museum Zoology"

ISSN 0968-0470 



Bulletin of 

The Natural History 

Museum 



I TUEMATUR W 

HISTORY MUSEUM 

11 DEC 1996 
PRESENTED 
I ^QOLOGYUBR a 



Zoology Series 




VOLUME 62 NUMBER 2 28 NOVEMBER 1996 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
(Natural History) ), instituted in 1949, is issued in four scientific series, Botany, 
Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Zoology Series is edited in the Museum's Department of Zoology 
Keeper of Zoology Dr C.R. Curds 

Editor of Bulletin: Dr N.R. Merrett 

Assistant Editor: Dr BT. Clarke 



Papers in the Bulletin are primarily the results of research carried out on the unique and ever- 
growing collections of the Museum, both by the scientific staff and by specialists from elsewhere 
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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) 
© The Natural History Museum, 1996 



Zoology Series 
ISSN 0968-0470 Vol. 62, No. 2, pp. 71-132 

The Natural History Museum 

Cromwell Road 

London SVV7 5BD Issued 28 November 1996 

Typeset by Ann Buchan (Typesetters), Middlesex 

Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 



Bull. not. Hist. Mas. Loud. (Zool.) 62(2): 71-82 



Issued 28 November 1996 



Indian Ocean echinoderms collected during 
the Sindbad Voyage (1980-81): 3. Ophiuroidea 
and Echinoidea 



ANDREW R.G. PRICE 

Ecosystems Analysis and Management Group, Department of Biological Sciences, University of Warwick, 
Coventry CV4 7AL, UK 



FRANCIS W.E. ROWE 

Goldbrook Boarding Kennels, Nuttery Vale, Cross Street, Hoxne, Suffolk IP21 5BB, UK 



W kAYURAi 

HISTORY MUSE! 

11 DEC 1996 
PRESENTED 



ZOOLOGY U BR> 



SYNOPSIS. At least 44 ophiuroid and 1 1 echinoid species are recorded from echinoderm collections made during an 
international expedition, the Sindbad Voyage, from Oman to China. Sampling localities include the little known Lakshadweep 
(Laccadive), Islands and Pula We (Sumatra) from which l\ c /c of the species were recorded. Following the zoogeographic 
subdivisions of Clark & Rowe ( 1971 ). range extensions are recorded for ten of theophiuroids: W. India (Amphioplus (Lymanella) 
sp.); Sri Lanka {Ophiactis modesta, Ophiarachna robillardi, Ophiodyscrita instratus); Maldives area (Cryptopelta granulifera, 
Ophiochaeta hirsuta); and Indonesia / East Indies (Amphiura (Amphiura) dejectoides. Amphiura (Amphiura) micra, Amphioplus 
{Amphioplus) stenaspis, Ophiogymna pellicula). In addition to the taxonomic treatment, ecological information for each 
echinoderm species (habitat types, depth range) is provided and broadly analysed. 



INTRODUCTION 



MATERIALS AND METHODS 



The systematics and distribution of Indian Ocean ophiuroids and 
echinoids are treated in detail by Clark & Rowe (1971). Regions for 
which limited information is available include the Lakshadweep 
(Laccadive) Islands, Sumatra and other parts of SE Asia. Recent 
studies including the systematics and zoogeography of ophiuroids 
and echinoids for the Lakshadweeps include Nagabhushanam & 
Rao (1972) and James (1989), the latter yielding many new species 
records for both echinoderm classes, which fill in gaps in the 
distribution records of Clark & Rowe (1971). Recent work has also 
been undertaken in SE Asia, including Indonesia (Aziz, 1981 ) and 
the west coast of Thailand (Bussarawit & Rowe, 1985; Bussarawit, 

in prep.). 

This paper reports on collections of ophiuroids and echinoids 
from these areas and other localities during an international, 
transdisciplinary voyage across the Indian Ocean from Oman to 
China. The expedition, Sindbad Voyage, was undertaken in 1 980-8 1 
aboard a replica of an ancient Arab sailing vessel, 'Sohar'. In 
addition to a systematic account, the zoogeographic significance of 
the results and the ecology of each species are broadly assessed. 
Details of the holothurian collections (Price & Reid, 1985) and 
asteroid collections (Marsh & Price, 1991 ) resulting from the expe- 
dition have already been published. Details of the crinoids collected 
are also being prepared (Marshall & Price, in prep.), and a detailed 
analysis of the ecology and biogeography of all five echinoderm 
classes will follow. 

Address for correspondence: First author 
© The Natural History Museum, 1996 



Specimens were collected by one of us (A. R.G. P.) and other expedi- 
tion members from localities at Muscat, Oman; Chetlat, Laks- 
hadweeps (Laccadives); SW India; SW Sri Lanka; and Pula We, 
Sumatra. Details of the sampling localities are shown in Figure 1. 
Sampling was undertaken principally on coral reefs using scuba. At 
each locality details of habitat type and depth range were recorded, 
along with the number of individuals of each species. The number of 
specimens collected is placed in parenthesis after each station 
number in the Material lists for each species. 

Material was fixed and preserved using standard methods (Lin- 
coln & Shields, 1979). Although several specimens (inadvertently 
included with the asteroid collections) had been identified earlier by 
L.M. Marsh and a few preliminary identifications were made in the 
field by A. R.G. P., specimens were mostly identified by and all 
species confirmed by F.W.E.R. who is also responsible for taxo- 
nomic comments. The ophiuroid and echinoid collections are 
deposited at the Natural History Museum, London, where the 
holothurian collection (Price & Reid, 1985) and a representative 
collection of the asteroids (Marsh & Price, 1991) have also been 
lodged. 

Following recent practice (e.g. Sloan, Clark & Taylor, 1979), 
systematic references are kept to a minimum by citing major works 
when possible (e.g. Clark & Rowe, 1971; Clark & Courtman Stock, 
1976; Cherbonnier & Guille, 1978; Rowe & Gates, 1995) from 
which the original species descriptions, recent authoritative diag- 
noses and taxonomic decisions can be traced. In some instances, 
further references are given (e.g. Sloan et ah, 1979) to provide 
additional systematic or biological information. 



72 



A.R.G. PRICE AND F.W.E. ROWE 



o 

50 E 



60 



70 



80 



90 



100 




Ug Bau 



Udjung 

Lo M« U 9 Ta Pa 




Fig. 1. (a) Map of northern Indian Ocean showing sampling areas (•) during Sindbad Voyage, with insert (b) for Pula We Sumatra. 



INDIAN OCEAN ECHINODERMS 



73 



RESULTS 



Class Ophiuroidea 



Family GORGONOCEPHALIDAE 

1. Astroboa nuda (Lyman, 1874) 

See. Clark &Rowe, 1971: 78; 92; Clark &Courtman Stock, 1976: 
108; 130; Baker, 1980: 60; Rowe & Gates, 1995: 364. 

Material. 8 1 0504D/3 ( 1 ), 8 1 0504D/4 ( 1 ). 

Collection sites. NW Klah, Pula We, Sumatra. 

Habitat and depth. Subtidal rock/coral; 13 m. 



Family AMPHIURIDAE 



2. Amphiura (Amphiura) dejectoides H.L. Clark, 1939 



See. 

33. 



Clark & Rowe, 1971: 80; 97; Cherbonnier & Guille, 1978: 



Material. 810501E/8 (4); 7810428D/5 (2). 

Collection sites. Ug Tapa Gadja & Ug Seukundo, Pula We, 
Sumatra. 

Habitat and depth. Subtidal rock / coral, coral reef; 10 & 15 m. 

Remarks. The specimens from stn 810501E/8 appear to accord 
well with the original description (Clark, 1939) of the species and 
descriptions in Clark & Rowe (1971) and Cherbonnier & Guille 
(1978). The 2 specimens from stn 810428D/5 differ in their firmer 
disc, with coarser ventral scaling and in having 6 arm spines 
proximally instead of fine ventral scaling and 5 proximal arm spines. 
They are identified as A. dejectoides with reservation. Clark (in 
Clark & Rowe, 1 97 1 ) suspected Amphiura inhacensis Balinsky may 
be conspecific with A. dejectoides H.L. Clark, a conclusion con- 
firmed by Cherbonnier & Guille ( 1978). The records included herein 
extend the distribution of this species eastwards across the Indian 
Ocean from the Red Sea, East Africa and Madagascar to the Indo- 
Malayan region. In the latter region it may prove to be widespread. 



3. Amphiura {Amphiura) micra H.L. Clark, 1938 

SEE. Clark & Rowe, 1971: 80; 97; Cherbonnier & Guille, 1978: 
46; Rowe & Gates, 1995: 350. 

Material. 810423B/2 (1). 

Collection sites. Ug Bau, Pula We, Sumatra. 

Habitat and depth. Coral, coral reef; 10-30 m. 

Remarks. This species is recorded across the tropical coast of 
Australia and from Madagascar. Its discovery at Pula We suggests a 
wider distribution in the Indo-Malayan region for this very small 
species. 



4. Amphiura (Amphichilus) ochroleuca (Brock, 1888) 

See. Clark & Rowe, 1971: 78; 100; Rowe & Gates, 1995: 344. 

Material. 810502C/1 (1), 810427B/3 (1). 

Collection SITES. SE Klah, SE Lho Pria Laot, Pula We, Sum- 
atra. 

Habitat and depth. In sponge and on subtidal rock; 0-10 m. 

Remarks. This species is known from the Indo-Malayan region 
and more or less circumscribes the Australian continental coastline 
(Rowe & Gates, 1995). Pula We appears to be the most westerly 
point of its distribution known to date. 

5. Amphioplus (Amphioplus) stenapsis H.L. Clark, 1938 

See. Clark & Rowe, 1971: 78; 101; Rowe & Gates, 1995: 344. 

Material. 810422B/4 (1). 

Collection sites. Nr. Klah / Seukundo, Pula We, Sumatra (disc 
only). 

Habitat and depth. Coral reef, 2-8 m. 

Remarks. Although the single specimen comprises only a com- 
plete disc with the bases of 2 arms (6 & 9 segments respectively), 
there is little doubt of its identity. This record extends the range of the 
species to Pula We from its type locality, Darwin, N Australia. The 
record of A. stenapsis from Madagascar by Cherbonnier & Guille 
(1978) is almost certainly based on a misidentification, judging by 
the very small size of the radial shields of their specimens. The 
confirmation of this species in the western Indian Ocean therefore 
requires confirmation. 

6. Amphioplus (Lymanella) sp. 

Material. 8 101 09 A/lb (2). 

COLLECTION sites. Beypore, India (west coast). 

Habitat and depth. Subtidal mud, 9 m. 

Remarks. Only the mouthparts and bases of the arms are avail- 
able to identify this taxon, which clearly represents a species of 
Amphioplus (Lymanella). The dorsal arm plates are trilobed, sug- 
gesting either species A. (L.) andreae (Liitken, 1872) or A. (L.) 
laevis (Lyman, 1874) in the key provided by Clark & Rowe (1971: 
102). Cherbonnier & Guille (1978) indicate that laevis has a wide 
range in the Indian Ocean and Indo-Malay region, whereas Clark & 
Rowe (1971) record andreae only from the Malay region. Without 
complete specimens it is not possible to determine the species nor 
indeed whether andreae and laevis are taxonomically separable. 



Family OPHIACTIDAE 

7. Ophiactis modesta Brock, 1 888 

See. Clark & Rowe, 1971: 105; Rowe & Gates, 1995: 379. 
Material. 8 10206 A/8 (3). 
Collection sites. Negombo, Sri Lanka. 
Habitat and depth. Coral/rock; 5 m. 



74 

Remarks. The 3 specimens were collected in a batch of 6 from 
stn 810206A/8 with 3 specimens of O. savignyi. They run 
unequivocably to the species O. modesta in the key provided by 
Clark & Rowe ( 1 97 1 : 1 05) where the relationship with other conge- 
ners is discussed by A.M. Clark (notes 25-27, pp. 103-105). Rowe 
(in Rowe & Gates, 1995) preferred to follow the views of Clark & 
Rowe (1971) regarding recognising the validity of the species O. 
modesta, a course also followed herein, rather than accept the 
sweeping synonymy of O. savignyi, which includes O. modesta, 
given by Cherbonnier & Guille (1978). 

8. Ophiactis savignyi (Miiller & Troschel, 1842) 

See. Clark & Rowe, 1971: 82; 103; Clark & Courtman Stock, 
1976: 164; Cherbonnier & Guille, 1978: 125; Sloan et ah, 1979: 
102; Price, 1983: 61; Rowe & Gates, 1995: 380. 

Material. 810206A/3 (5), 810206A/8 (3), 810426A/2 (1), 
810501E/1 (1), 810501G/3 (1), 8105011/2 (1), 810502F/4 (2). 

COLLECTION sites. Negombo, Sri Lanka; W Rubiah, Ug 
Seukundo, E Klah, Pula We, Sumatra. 

Habitat and depth. Sponge, coral/rock, subtidal rock, coral reef; 
5-14 m. 

Remarks. The specimens identified here are fissiparous and 6- 
armed with trilobed dorsal arm plates and proximally 6 arm spines, 
according well with the key characters given in Clark & Rowe 
(1971). 



Family OPHIOTRICHIDAE 

9. Gymnolophus obscura (Ljungman, 1867) 

See. Clark & Rowe, 1971: 82; 1 17; Rowe & Gates, 1995:411. 

Material. 810124A/11 (1), 810125A/2 (1), 810430A/20d (2). 

Collection sites. Ala Gala & Deumba Gala, Galle, Sri Lanka; 
Ug Seukundo, Pula We, Sumatra. 

Habitat and depth. Coral reef and epizoic on crinoids on subtidal 
rock; 8-15 m. 

Remarks. This species is commonly epizoic on comasterid 
crinoids, the host species for which have not been identified for the 
specimens recorded herein. 

10. Ophiothela danae Verrill, 1 869 

SEE. Clark & Rowe, 1971: 84; 116; Clark & Courtman Stock, 
1976: 141; Price, 1983: 63; Rowe & Gates, 1995:419. 

Material. 810125B/1 (3), 810206A/3/78 (4), 810425F/8 (10+), 
8 10428 A/2 (7), 810428A/14(3), 810501A/3 (30+), 810501A/4(1), 
810428B/1 (2), 810428D/3c (3), 810501E/9 (15+). 

Collection sites. Ala Galla, Galle & Negombo, Sri Lanka; N. 
Udjung Lo Me (NE Sabang Bay), Ug Bau, Ug Seukundo, Ug Tapa 
Gadja, Pula We, Sumatra 

Habitat and depth. Epizoic on macroalgae, gorgonian, fire coral 
(Millepora sp.) sponge and on holothurians (Thelenota ananas), all 
on rock /coral; 2-30 m. 



A.R.G. PRICE AND F.W.E. ROWE 

11. Ophiothrix exigua Lyman, 1 874 

See. Clark & Rowe, 1971: 84; 1 10; Cherbonnier & Guille, 1978: 
140; Rowe & Gates, 1995: 422. 

Material. 810206A/8 (5), 810502C/1 (2), 810502E/2 (3). 

Collection sites. Negombo, Sri Lanka; E Klah, Pula We, 
Sumatra. 

Habitat and depth. Coral/rock, sponge on coral reef and subtidal 
rock; 5-10 m. 

Remarks. One specimen from stn 810206A/8, both from 
810502C/1 and two from 810502E/3 are very juvenile specimens. 

12. Ophiothrix savignyi (Miiller & Troschel, 1842) 

See. Clark & Rowe, 1971: 84; 109; Cherbonnier & Guille, 1978: 
142; Price, 1983:65. 

Material. 801114B/1 (1). 

Collection sites. Muscat, Oman (1 specimen). 

Habitat and depth. Coral reef; 2 m. 

13. Ophiothrix trilineata Liitken, 1 869 

See. Clark & Rowe, 1971: 84; 111; Clark & Courtman Stock, 
1976: 145; Sloan etal, 1978: 103; Rowe & Gates, 1995: 423. 

Material. 810420A/5 (2), 810422B/3 (3), 810426A/2 (1), 
810428A/9 (2), 810428C/7 (1), 810428D/3a&b (6&2), 810430A/8 
(1), 810430A/18 (1), 810430A/21c (1), 810501D/2 (1), 810501E/2 
(3), 810501E/5 (1), 810501E/8 (1), 810501E/9 (2), 810501E/1 1 (1), 
810502F/2 (1), 810501F/3 (1), 810501G/1 (2), 810501G/3 (1 & 1 
juvenile), 810501G/5 (3), 810502C/2 (1 very juvenile), 810502D/4 
(4), 810502E/3 (4 juvenile), 810502F/2 (2 juvenile), 810502G/3 (2 
juvenile). 

Collection sites. Klah / E Klah, Nr. Seukundo, Ug Seukundo, 
Pula We. Sumatra, W Rubiah, Ug Bau, Ug Tapa Gadja, Pula We, 
Sumatra. 

Habitat and depth. Coral reef, soft coral / gorgonian, subtidal 
rock, sponge/subtidal rock, sponge; 2-30 m. 

Remarks. The majority of specimens exhibit the characteristic 
arm colour pattern of 5 longitudinal lines alternating white and dark 
blue. However, a few specimens bear a wide median pale longitudi- 
nal line along the arms, and at least one specimen (810501E/2) is 
distinctively patterned with cream blotches, the linear pattern being 
discernable only near the ends of the arms (see Clark & Rowe, 197 1 : 
111). Next to O. (A.) purpurea, this is the most common ophiuroid 
species collected. 

14. Ophiothrix (Acanthophiothrix) armata Koehler, 
1905 

SEE. Clark & Rowe, 1971: 84; 1 1 1; Rowe & Gates, 1995: 423. 

Material. 810422B/4 (15+), 810502F/6 (2). 

Collection sites. Klah / Nr. Seukundo, Pula We, Sumatra. 

Habitat and depth. Coral reef, 2-8 m. 

Remarks. This species is recorded from the Indo-Malay region, 
tropica] Australian coasts and the South Pacific (Clark & Rowe, 



INDIAN OCEAN ECHINODERMS 



75 



197 1 ). The present record is the most westerly so far known for the 
species. 

15. Ophiothrix (Acanthophiothrix) purpurea von 

Martens, 1867 

See. Clark & Rowe, 1971: 86; 1 12; Cherbonnier & Guille, 1978: 
148; Sloan etal., 1978: 103; Rowe & Gates, 1995: 423. 



Material. 810204A/4 (3), 8 10421 A/2 (1), 8 10421 A/9 (1 juve- 
nile), 810421B/1 (1), 810422D/3 (2), 8 10423 A/4 (1), 810423B/2 
(1), 810425D/2(1), 80425D/4b(l), 810425F/7 (7), 810425F/8 (1), 
810426A/2 (3), 8 10427 A/2 (2), 810428A/2 (2), 810428A/7 (3), 
810428A/8 (4), 810428A/9 (2), 810428D/3f (23), 810430A/3 (1), 
810430A/21c (6), 810430A/22b (1), 810430A/26 (5), 8 10501 A/ 1 
(1), 8 10501 A/2 (1 juvenile), 8 10501 A/4 (1 juvenile), 8 10501 A/6 
(9), 810501E/7 (2), 810501E/9 (1), 81050IE/14 (1), 810501E/15 
(1), 810504B/2 (1). 

Collection sites. SW Kalpitiya, Sri Lanka; Ug Bau, Rubiah, Ug 
Seukundo, UgTapa Gadja, N Klah, ?N. Udjung Lo Me, NE Sabang 
Bay, Pula We, Sumatra. 

Habitat and depth. Coral reef, soft coral, fire coral (Millepora 
sp.), subtidal rock/coral (epizoic on soft coral / gorgonian & crinoid, 
sponge); 2-30 m. 

Remarks. The most common ophiuroid species collected. 



16. Ophiothrix (Acanthophiothrix) spinosissima 

Koehler, 1905 

SEE. Clark & Rowe, 1 97 1 : 86; 1 1 2. 

Material. 810422B/4 (4), 810501G/5 (2), 810502F/6 (1). 

Collection sites. Ug Seukundo, Klah / Ug Seukundo, Pula We, 
Sumatra. 

Habitat and depth. Coral reef, 2-8 m. 

Remarks. The specimens run down well to O. (A.) spinossima in 
the key provided by Clark & Rowe (1971). However, 2 specimens 
from stn 8 10422B/4 have a single dark line running the length of the 
dorsal side of the arms, rather than a series of dark spots. 

17. Macrophiothrix aspidota (Miiller & Troschel, 1842) 

SEE. Clark, 1968: 285; Clark & Rowe, 1971: 114; Clark & 
Courtman Stock, 1976: 137; Hoggett, 1992: 91. 

Material. 810123B/6 (1), 810124A/8 (1), 810206A/6 (1), 
810206A/7 (1). 

Collection sites. Negombo, Closenburg Point, Galle. Ala Gala, 
Galle, Sri Lanka. 

Habitat and depth. Subtidal rock, coral / rock; 5-15 m. 



18. Macrophiothrix demessa (Lyman, 1861) 

See. Clark, 1968: 289; Clark & Rowe, 1971: 82; 1 14; Hoggett, 
1991: 1089; 1992: 1 17; Rowe & Gates, 1995:412. 

Material. 801212B/1 (1), 810424B/4 (2). 

Collection sites. Chetlat, Lakshadweep (Laccadive) Islands; 
Seulakoe, Pula We, Sumatra. 



Habitat and depth. Coral rubble, coral reef; 8 m & 20-30 m. 

Remarks. A.M. Clark ( 1 968) transferred this species to the genus 
Macrophiothrix referring Ophiothrix (Amphiophiothrix) H.L. Clark, 
1946, of which demessa is type (and only) species to the synonymy 
of Macrophiothrix H.L. Clark, 1938. 

19. Macrophiothrix elongata (H.L. Clark, 1938) 

See. Clark, 1968: 291; Clark & Rowe, 1971: 82; 114;Price, 1983: 
61; Hoggett, 1992: 125. 

Material. 801 1 1 1 A/5 ( 1 ), 801 1 14A/5 (2), 801 1 14B/2 (1 ). 

Collection sites. Muscat harbour, Oman. 

Habitat and depth. Coral reef, subtidal rock/coral/sand; 0.5-3 
m. 

20. Macrophiothrix longipeda (Lamarck, 1816) 

See. Clark, 1968: 300; Clark & Rowe, 1971: 82; 1 14; Clark & 
Courtman Stock, 1976: 139; Hoggett, 1991: 1103; Hogett, 1992: 
151; Rowe & Gates, 1995:413. 

Material. 80 1 2 1 2B/3 ( 1 ), 8 1 0206A/5 ( 1 ), 8 1 0430A/24a ( 1 ). 

Collection SITES. Chetlat, Lakshadweep (Laccadive) Islands; 
Negombo, Sri Lanka; Ug Seukundo, Pula We, Sumatra. 

Habitat and depth. Coral/rock, coral rubble; 5-10 m. 

21. Macrophiothrix lorioli A.M. Clark, 1968 

See. Clark, 1968: 302; Clark & Rowe, 1971: 82; 115; Hoggett, 
1991: 1108; Hoggett, 1992: 161; Rowe & Gates, 1995:414. 

Material. 810502D/2 (1), 810502E/1 (1). 

Collection sites. E Klah, Pula We, Sumatra. 

Habitat and depth. Coral reef, coral reef/subtidal rock; 5 & 
10 m. 

22. Macrophiothrix nereidina (Lamarck, 1816) 

SEE. Clark & Rowe, 1971: 86; 107 (as Ophiothrix (Keystonea) 
nereidina); Hoggett, 1992: 228 (as Macrophiothrix); Rowe & Gates, 
1995: 426 (as O. (Keystonea) nereidina). 

Material. 8 10421 A/4 (1), 810422B/3 (2), 810430A/20d (1), 
8 1 0430 A/2 lc (2), 8 1 0430A/22b ( 1 ), 810501E/6 (1). 

Collection sites. Ug Seukundo, Pula We, Sumatra. 

Habitat and depth. Coral reef, coral / rock; 2-10 m. 

REMARKS. The species nereidina (which was placed in the 
subgenus Ophiothrix (Keystonea) by A.M. Clark, 1967) is included 
herein in the genus Macrophiothrix. Hoggett (1991) stated that 'it is 
particularly difficult to determine the respective boundaries between 
Macrophiothrix H.L. Clark, 1938 and two subgenera of Ophiothrix, 
O. (Placophiothrix) H.L. Clark, 1938 and O. (Keystonea) A.M. 
Clark, 1967.' The differences between these taxa have traditionally 
relied principally on arm length, shape of dorsal arm plates, relative 
spinular armament of the disc plates including cover of the radial 
plates (see A.M. Clark, 1967; Clark & Rowe, 1971). The difficulty 
in recognising the supraspecific limits of these taxa is made all the 
more obvious by the treatment of the species Macrophiothrix 



76 



A.R.G. PRICE AND F.W.E. ROWE 



propinqua, placed in the subgenus Keystonea by A.M.Clark (1967), 
whilst Devaney (1974) described O. (Placophiothrix) westwardi 
which has been considered conspecific with propinqua by Hoggett 
(1991). Later, in a far-reaching and critical treatment of Macro- 
phiothrix, Hoggett (1992: PhD thesis) commits both the taxa 
Placophthiothrix and Keystonea to the synonymy of Macrophio- 
thrix, transferring the included species of the former two taxa to the 
latter taxon and to whom this move is herein credited. 

23. Macrophiothrix propinqua (Lyman, 1861) 

See. Clark & Rowe, 1971: 86; 107 (as Ophiothrix {Keystonea) 
propinqua); Clark, 1980: 537; Hoggett, 1991: 1 130; Hoggett, 1992: 
204; Rowe & Gates, 1995: 415. 

Material. 810428D/5 (5), 810501C/3 (1 juvenile), 810501D/1 
(1 juvenile), 810501E/13 (1), 810501DF/1 (1 juvenile), 810501G/1 
(1), 810501G/5 (3), 8105011/2 (2), 810502F/3 (1), 810502G/3 (1). 

Collection sites. Ug Tapa Gadja, Ug Seukundo, E Klah, Pula 
We, Sumatra. 

Habitat and depth. Subtidal rock/coral, coral rubble, coral reef, 
soft coral; 2-20 m. 

Remarks. This species was first transferred to the genus 
Macrophiothrix H.L. Clark, 1938 by A.M. Clark (1980) from 
Ophiothrix (Keystonea) A.M. Clark, 1967. 

24. Macrophiothrix variabilis (Duncan, 1887) 

See. Clark, 1968: 308; Clark & Rowe, 1971 : 1 15; Hoggett, 1991: 
1138; Hoggett, 1992: 218; Rowe & Gates, 1995: 416. 

Material. 810206A/4 (1). 

Collection sites. Negombo, Sri Lanka. 

Habitat and depth. Coral / rock, 5 m. 

25. Macrophiothrix virgata (Lyman, 1861) 

See. Clark & Rowe, 1971: 86; 113 (as Ophiothrix (Placophio- 
thrix) virgata); Hoggett, 1992: 236. 

Material. 810423D/2 (1). 

Collection sites. Ug Bau, Pula We, Sumatra. 

Habitat and depth. Coral reef; 2-8 m. 

REMARKS. See remarks under Macrophiothrix nereidina. 

26. Ophiogymna pellicula (Duncan, 1876) 

See. Clark & Rowe, 1971: 84; 117; Clark & Courtman Stock, 
1976; 140 (as O.fulgens); Rowe & Gates, 1995: 417. 

Material. 810504B/2 (1). 

Collection sites. Rubiah, Pula We, Sumatra. 

Habitat and depth. Soft coral; 10 m. 

Remarks. The single specimen has a d.d. = 2.7 mm, a.l. = c. 20 
mm. The disc is mottled cream and pink, and the arms are banded 
with wide pink and narrow cream bands. An irregular longitudinal 
line of cream spots is evident along the dorsal midline of the arms. 
The disc is covered (except for the radial shields) with minute, 



pointed granules with larger, conical spines interradially towards the 
edge of the disc. Clark & Courtman Stock (1976) include 
Placophiothrix phrixa H.L. Clark as a synonym of O. fulgens 
(Koehler) which in turn is included in the synonymy of O. pellicula 
by Rowe (in Rowe & Gates, 1995). The species therefore appears to 
to be distributed from the Gulf of Aden to the Indo-Malay region and 
the NW coast of Australia in depths of 10-1 16 m. 

27. Ophiopteron elegans Ludwig, 1888 

See. Clark & Rowe, 1971: 84; 115; Rowe & Gates, 1995: 419. 
Material. 810428D/3e (2), 810428D/5 (8). 
Collection sites. Ug Tapa Gadja, Pula We, Sumatra. 
Habitat and depth. Coral reef, subtidal rock/coral; 15 m. 



Family OPHIOCOMIDAE 

28. Ophiarthrum pictum Miiller & Troschel, 1842 

See. Clark & Rowe, 1971: 86; 121; Rowe & Gates, 1995: 385. 

Material. 810502H/1 (1). 

Collection sites. E Klah, Pula We, Sumatra. 

Habitat and depth. Subtidal rock, 2m. 

29. Ophiocomella sexradia (Duncan, 1887) 

See. Clark & Rowe, 1971: 86; 118; Devaney, 1974: 162; 
Cherbonnier & Guille, 1978: 179; Rowe & Gates, 1995: 389. 

Material. 81042D8/3d (1). 

Collection sites. Ug Tapa Gadja, Pula We, Sumatra. 

Habitat and depth. Coral reef; 15 m. 

Remarks. The single specimen from Pula We measures d.d. = 3 
mm; a.l. = 10+ mm (broken near tip). With the exception that the 
uppermost of the 4 arm spines is distinctly longer than the lower 
ones, the character separating Ophiomastix sexradiata A.H. Clark 
1952 (known only from its type locality: Bikini Atoll, Marshall Is, 
SW Pacific) from Ophiocomella sexradia (Duncan)(identified 
throughout the Indo-West Pacific region and possibly tropicopolitan) 
in Clark & Rowe's (1971) key, all other skeletal characters of the 
present specimen accord with those described as fitting O. sexradia 
(note 65, p. 118) by A.M. Clark. Cherbonnier & Guille (1978), 
following their study of Malagasy material, concur with the com- 
ments expressed by A.M. Clark (in Clark & Rowe, 1971) and 
tentatively consider Ophiomastix sexradiata to be conspecific with 
Ophiocomella sexradia simultaneously agreeing with A.M. Clark 
that the status of the genus Ophiocomella as distinct from Ophio- 
coma remains doubtful. Although the present specimen does nothing 
to clarify the generic status of Ophiocomella, it does support strongly 
the view that the two nominal species are conspecific and confirm 
the synonymy proposed by Cherbonnier & Guille (1978). 

30. Ophiocoma dentata Miiller & Troschel, 1 842 

See. Devaney, 1970: 13; Clark & Rowe, 1971: 86; 1 19; Rowe & 
Gates, 1995: 386. 

Material. 810502H/2b (1). 



INDIAN OCEAN ECHINODERMS 



77 



Collection sites. E Klah, Pula We, Sumatra. 

Habitat and depth. Subtidal rock, 2m. 

Remarks. This single specimen (d.d. c. 11.3 mm, distorted) 
exhibits two of the described colour forms for the species. At the 
centre of the dorsal surface of the disc is a cream spot (c. 1.1 mm 
diameter). This is surrounded by an irregular ring (c. 2.2 mm wide) 
which is uniformly dusky/grey with darker spots. The remainder of 
the disc dorsally and ventrally is reticulated dusky/grey on a cream 
background. 

31. Ophiocoma erinaceus Miiller & Troschel, 1842 

SEE. Clark & Rowe, 1971: 1 14; 1 1 9; Clark & Courtman Stock, 
1976: 173; Sloan era/, 1979: 106; Bussarawit & Rowe, 1985: 1 (as 
O. similanensis n. sp.); Rowe & Gates, 1995: 387. 

Material. 801212A/3 (1), 810422B/3 (10), 810427B/1 (1), 
810427D/5 (1), 810428C/6 (2 juveniles), 810428E/5 (1 juvenile), 
810430A/3 (2), 810501F/1 (1), 810501G/5 (3), 810501G/6 (2), 
810501K71 (2). 

Collection sites. Chetlat, Lakshadweep (Laccadive) Islands; 
Ug Seukundo, Ug Bau, Lho Pria Laot, Ug Murung, Ug Tapa Gadja, 
Pula We, Sumatra. 

Habitat and depth. Coral reef, coral conglomerate, coral rubble, 
subtidal rock; 2-25 m. 

Remarks. This is the commonest species of Ophiocoma col- 
lected. The collection comprises some 26 specimens, ranging in size 
from d.d. = 3.6-22.2 mm, which show clearly both colour changes 
and development of disc granulation with growth. Juveniles up to 
d.d. = c. 5 mm bear no granules and are usually marked radially 
across each radial shield with a cream line, as described by Bussarawit 
& Rowe (1985) for their new species O. similanensis. By d.d. = 5.7 
mm granules are developed at the centre of the disc and along 10 
radiating lines to the edge of the disc where an irregular line of 
granules is developed around the periphery joining these radiating 
lines and thus leaving bare the dorsal interradial and radial portions 
of the disc. By d.d. = c. 1 1 mm granules are developed over the 
interradial but not radial regions of the disc, but granules are still not 
developed ventrally. By d.d. = 12.5 mm granules cover the whole 
surface of the disc except for the radial shields which remain bare, 
while granules begin to extend in a wedge shape, on the ventral side 
of the disc. This arrangement may remain in specimens up to d.d. = 
14 mm but generally from about d.d. = > 13 mm the radial shields 
become covered by granules. In specimens up to d.d. = 12.5 mm 
some central and peripheral granules may be more prominent by 
their slightly more elongate shape, but in larger specimens granules 
are more evenly rounded and more or less evenly sized. The cream 
colour pattern disappears with increased d.d. and is absent in speci- 
mens with d.d. > 1 1 mm. 

It is very clear that Ophiocoma similanensis Bussarawit & Rowe, 
1985 is based on juvenile specimens of O. erinaceus, to the syn- 
onymy of which O. similanensis is herein committed. 

32. Ophiocoma pica Miiller & Troschel 

See. Devaney, 1970: 25; Clark & Rowe, 1971: 86; 1 18; Clark & 
Courtman Stock, 1976: 173; Sloan etal, 1979: 106; Rowe & Gates, 
1995: 387. 

Material. 8012I2A/3 (1). 

Collection sites. Chetlat, Lakshadweep (Laccadive) islands. 



Habitat and depth. Coral reef; 20 m. 

33. Ophiocoma pusilla (Brock, 1888) 

See. Devaney, 1970: 25; Clark & Rowe, 1971: 86; 118; Clark & 
Courtman Stock, 1976: 174; Sloan era/, 1979: 106; Rowe & Gates, 
1995: 388. 

Material. 810422E/4 (1), 810425C/2 (1), 810430A/21c (1), 
810501E/3 (1), 810501K/4 (1). 

Collection sites. N Klah island, Sabang Bay, Ug Seukundo, 
Pula We, Sumatra. 

Habitat and depth. Coral/sand, coral conglomerate, coral reef; 
2-10 m. 

Remarks. The 5 specimens range in size from d.d. = 3-7.5 mm. 
The characteristic, enlarged, tissue-covered arm spines (see Clark & 
Rowe, 1971 ) appear on specimens from d.d. > 5 mm. 

34. Ophiomastix annulosa (Lamarck, 1816) 

SEE. Clark & Rowe, 1971: 86; Rowe & Gates, 1995: 390. 

Material. 810123A/3 (1), 810212A/3 (2), 810213A/4 (1). 

COLLECTION sites. Kakoni rocks, Pigeon Island & Unawatuna, 
Galle, Tangalla, Sri Lanka. 

Habitat and depth. Coral reef, subtidal rock/coral, subtidal 
rock; 3-10 m. 

35. Ophiomastix caryophyllata Liitken, 1869 

See. Clark&Rowe, 1971: 86; 120; Cherbonnier & Guille, 1978: 
185; Rowe & Gates, 1995: 390. 

Material. 810425C/1 (2), 810428C/7 (7 2 ), 810430A/21b (2). 

COLLECTION sites. E Sabang Bay, Ug Bau, Ug Seukundo, Pula 
We, Sumatra 

Habitat and depth. Coral reef, coral/rock; 3-10 m. 



Family OPHIONEREIDAE 

36. Ophionereis dubia (Miiller & Troschel, 1842) 

See. Clark & Rowe, 1971: 122; Clark & Courtman, 1976: 179; 
Price, 1983: 67; Rowe & Gates, 1995: 408. 

Material. 810502D/5 (1). 

Collection sites. E Klah, Pula We, Sumatra. 

Habitat and depth. Subtidal rock/sand; 10 m. 

37. Ophionereis fusca Brock, 1888 

See. Clark, A.M. 1953: 69; 78; Clark & Rowe, 1971: 88; 122; 
Rowe & Gates, 1995:408. 

Material. 810421B/2 (1). 

Collection sites. Nr. Seukundo, Pula We, Sumatra. 

Habitat and depth. Subtidal sand; 10-20 m. 



78 



A.R.G. PRICE AND F.W.E. ROWE 



Family OPHIODERMATIDAE 

38. Cryptopelta granulifera H.L. Clark, 1909 

See. Clark & Rowe, 1971: 88; 128; Rowe & Gates, 1995: 394. 

Material. 801212B/1 (1). 

Collection sites. Chetlat, Lakshadweep (Laccadive) Islands. 

Habitat and depth. Coral rubble; 8 m. 

Remarks. Originally described from Mauritius. Rowe & Gates 
(1995) describe the distribution as including tropical Australia, the 
Indo-Malayan region and Philippine Islands. The specimen is iden- 
tified here from the Laccadive Islands for the first time. 

39. Ophiarachna affinis Liitken, 1 869 



occurrence of spinelets on the discs of 4 specimens from Aldabra, 
western Indian Ocean and 2 specimens from Palau in the western 
Pacific Ocean which they examined suggested that Ophiochaeta 
boschmai A.H. Clark, 1964 is a synonym of Ophiochaeta hirsuta 
Liitken, 1869. They pointed out that Cherbonnier & Guille,1978, 
had described a new species, O. crinita, based on a single specimen, 
from Madagascar, but did not comment further. In the present 
collection the specimen from the Lakshadweep Islands accords with 
the description of boschmai in that the disc is granule-covered 
dorsally but bears spinelets on its ventral surface. The specimen 
from Pula We, on the other hand, accords with the description of 
crinita in that the disc is covered dorsally and ventrally by elongate 
spines. Considering the comments by Sloan et al. (1979), the two 
specimens reported herein are referrred to O. hirsuta, with the 
implication that O.crinita Cherbonnier & Guille should also be 
referred to the synonymy on the basis that it exhibits the extreme 
spiny form of O. hirsuta. 



I; 123; Sloan era/., 1979: 111; Rowe 42. Ophiodyscrita instratus (Murakami, 1944) n. comb. 



SEE. Clark & Rowe, 1971: 
& Gates, 1995: 395. 

Material. 810425C/la,b (1). 

Collection sites. E. Sabang Bay, Pula We, Sumatra. 

Habitat and depth. Coral conglomerate; 3-6 m. 

Remarks. A.M. Clark (in Clark & Rowe, 1 97 1 : Note 83, p. 1 23) 
describes in detail colour variation in Ophiarachna affinis and O. 
mauritiensis de Loriol, concluding that specific distinction between 
the two is difficult to make. In the present specimens with d.d. = 22 
mm, the disc is uniformly brownish-grey and the dorsal side of the 
arms has a broad longitudinal central dusky band either side of 
which is a narrower pale band, the 3 bands being demarcated by 4 
irregular, very narrow longitudinal dark bands. This corresponds to 
A.M. Clark's form C colour pattern. If O. affinis and O. mauriti- 
ensis are conspecific the species is clearly widely distributed in the 
Indo-West Pacific region. 

40. Ophiarachna robillardi de Loriol, 1893 



See. Clark & Rowe, 1971: 



123. 



Material. 810126B/4 (1), 810213A/3 (2). 

Collection sites. Galle, Tangalla, Sri Lanka. 

Habitat and depth. Coral reef, 3-5 m. 

Remarks. This is a significant extension of range for this species 
described from Mauritius. The species is recorded as having 5 arm 
spines (H.L. Clark, 1909; size not recorded) but the present 3 
specimens have 7-9 arm spines at d.d. = 21.5 mm; 9-10 arm spines 
at d.d. 31.5 mm and 10-1 1 arm spines at d.d. = 36.5 mm. 

41. Ophiochaeta hirsuta Liitken, 1869 

See. Clark&Rowe, 1971:88; 127; Sloan et al, 1979: 115;Rowe 
& Gates, 1995 398. 

Material. 801212B/1 (1), 810425C/2 (1). 

Collection sites. Chetlat, Lakshadweep (Laccadive) Islands; E. 
Sabang Bay, Pula We, Sumatra. 

Habitat and depth. Coral rubble, 3-8 m. 

Remarks. Sloan et al. (1979) concluded that variation in the 



See. Murakami, 1944: 272 (as Ophiostegastus instratus): 
A.M.Clark, 1968: 320 (as Ophiostegastus instratus; discussion) 

Material. 810124A/4 (1). 

Collection sites. Ala Gala, Galle, Sri Lanka. 

Habitat and depth. Subtidal rock, 10-15 m. 

Remarks. This specimen, apart from its smaller size and fewer 
naked disc plates, accords so well with Murakami's (1944) descrip- 
tion of Ophiostegastus instratus that its identity is in no doubt. The 
species is, however, transferred to the genus Ophiodyscrita H.L. 
Clark, 1938 (type-species O. acosmeta H.L. Clark), with 
Ophiostegatus Murakami, 1944 (of which instratus is the type- 
species) reduced to a junior synonym of Ophiodyscrita. The 
distinctness of two genera has been questioned by A.M. Clark 
(1968) and Guille & Vadon (1985) on the grounds of variation of 
granulation with increased specimen size. Tabulation of measure- 
ments (Table 1 ) taken from original species descriptions and similar 
details of the specimens from Sri Lanka, shows an interesting 
picture. From this table it becomes apparent that only two species 
can be recognised: a) Ophiodyscrita instratus (Murakami, 1 943)(d.d. 
= 7 mm) of which the larger Ophiostegastus novaecaledoniae Guille 
& Vadon (d.d. = 9-1 1 mm) is a synonym, being an extreme form of 
instratus in which many disc plates, including the radial shields, 
have become prominent (convex) and naked of granules; this species 
possesses supplementary oral shields (granule covered in small 
specimens < 7 mm d.d.) and b) O. acosmeta H.L. Clark (d.d. - 5 
mm) with which O. pacifica (Murakami, 1943)(d.d. 4 mm) and 
Ophiostegastus compsus A.M. Clark (1968) type locality Bahrain 
(d.d. = 8-10.5 mm) appear to be conspecific. O. acosmeta has an 
even covering of granules over the disc which are gradually lost only 
from the oral shields (as in the type of acosmeta; d.d. = 5 mm) and 
adoral plates (as in the type series of compsus; d.d. = 8-10.5 mm; 
though A.M. Clark (1968) does note that the largest paratype of 
compsus (d.d. = 10.5 mm) has a small bare patch dorsally at the base 
of 4 of the arms). Supplementary oral shields are absent from 
compsus, according to A.M. Clark, and are not recorded for either 
acosmeta or pacifica. In both instratus and acosmeta, as recognised 
herein, it is clear that arm spine number increases with size. 

There is clearly insufficient justification for recognising 
Ophiodyscrita and Ophiostegastus as, separate genera on the basis of 
the extent of disc granulation, for it is clear (see Clark & Rowe, 
1971) that such differences occur between species included within 



INDIAN OCEAN ECHINODERMS 



79 



Table 1. 


Details of species Ophiodyscrita acosmeta H.L. Clark* and O. instratus (Murakami)* 


Taxon 


d.d. (mm) a.sp. l.a.sp. s.o.s. Granulation (disc) 



Ophiocryptus* 


4.00 


6-5 


1/3 seg. 


_ T 


pacificus Murakami, 1943 










Ophiodyscrita* 


5.0 


8(7) 


1/2 seg. 


_ 7 


acosmeta H.L. Clark, 1938 










Sindbad spec* 


5.3 


6-5 


<l/2 seg. 


+ (granule 
covered 


Ophiostegastus* 


7.00 


7 


1/3 seg. 


+ (naked) 


instratus Murakami, 1944 










Ophiostegastus* 


8-10.5 


9 


< 1/2 seg. 


- 


compsus A.M. Clark, 1968 










Ophiostegastus* 


9-11 


9-10 


< 1/2 seg. 


+ (naked) 


novaecaledoniae 










Guille & Vadon, 1985 











Complete cover dorsal and ventral 

Complete cover dorsal and ventral except 2 oral shields (Clark & Rowe 

(1971: 135) 
Complete cover dorsal and ventral except radial plate at base of deach arm. 

each mid marginal plate and ventrally, each of the oral shields 
Complete except 3 plates at base of each arm; each mid marginal; each oral 

shield and supplementary oral shield 
Complete except each oral shield and adoral shield (A.M. Clark notes a 

small bare patch as base of 4 arms of one paratype d.d. = 10.5 mm) 
Many dorsal plates, including radial shields, marginal plates and ventral 

plates including oral, supplementary oral and adoral shields bare of 

granules 



d.d. = disc diameter; a.sp. = number of arm spines; l.a.sp. = length of arm spines; s.o.s. (+/-) = presence/absence of supplementary oral shields. 



the recognised limits of other ophiodermatid genera (e.g. Ophiopeza 
& Ophiarachnella). Similarly, the occurrence of supplementary oral 
shields is also a variable character. The recognition of Ophiodyscrita 
within the family appears, therefore, to rely, more or less solely, on 
the extension of granulation along the arms. The genus is closely 
related to Ophiopeza with which it shares the possession of a triangle 
of 3 plates between the radial shields (see Vail & Rowe, 1989). The 
record of O. instratus from Sri Lanka greatly extends the known 
distribution of the species from Japan and New Caledonia (S.W. 
Pacific). The distribution of the genus, Ophiodyscrita (syn: 
Ophiostegastus ) is clearly widespread in the Indo-West Pacific 
region. 

43. Ophiopsammus yoldii (LUtken, 1856) 

| See. A.M.Clark, 1968; 317; Clark & Rowe, 1971:90; 127;Vail& 
, Rowe, 1989: 277; Rowe & Gates. 1995: 402. 

Material. 810430A/22b (1). 

I Collection SITES. Ug Seukundo, Pula We, Sumatra. 

Habitat and depth. Coral rubble; 9 m. 

Remarks. Nearly half of the disc of this small specimen has been 
lost leaving the remaining disc (d.d. = c. 6 mm) and three of the 
original arms, which are also damaged (a.l. = c. 1 7 mm; d.d./a.l. = c. 
3+ : 1). Arising from the damaged edge of the disc are three new, 
minute arms at slightly different stages of growth judging by the 
relative development of the ventral arm plates on each arm. These 
arms are also damaged but the longest is judged to have been not 
more than c. 3-4 mm in length. Following Vail & Rowe's (1989) 
revision of the genus Ophiopsammus, there is no reason for not 
identifying the specimen from Pula We as O. yoldii, for it appears to 
match their criteria for the species even though it is of small size. The 
species is not known to be fissiparous. Although this may be the first 
observation of fissiparity in O. yoldii, the development of six arms, 
in this case, may be an unusual response to severe damage, rather 
than being related to an asexual reproductive strategy. This matter 
requires further investigation. 

Family OPHIURIDAE 

44. Ophiolepis cincta Miiller & Troschel, 1842 

See. Clark & Rowe, 1971: 90; 129; Clark & Courtman Stock, 



1976: 189; Sloan etal, 1979: 1 15-1 17; Rowe & Gates, 1995: 399. 
Material. 801212B/1 (1). 

Collection SITES. Chetlat, Lakshadweep (Laccadive) Islands. 
Habitat and depth. Coral rubble; 8 m. 

Class Echinoidea 

Family CIDARIDAE 

1. Eucidaris metularia (Lamarck, 1816) 

See. Clark & Rowe, 1971: 140; 150; Clark & Courtman Stock, 
1976: 215; Sloan etal, 1979: 1 1 7 Rowe & Gates, 1995: 195. 

Material. 810426A/4 (1), 810501E/12 (1). 

Collection sites. Rubiah, Ug Seukundo, Pula We, Sumatra. 

Habitat and depth. Coral reef, coral aggregate; 10 & 14 m. 



Family DIADEMATIDAE 

2. Diadema setosum (Leske, 1778) 

See. Clark & Rowe, 1971: 140; 153; Clark & Courtman Stock, 
1976: 226; Sloan etal., 1979: 118; Price, 1983: 73; Rowe & Gates, 
1995: 207. 

Material. 801027A/1 (1), 801030A/la,b (4), 810212A/1 (1), 
8 1 0420 A/ 1 (1), 810420A/4 (1), 8 10426 A/3 (1), 810426B/5 (2), 
810426B/12 (1), 810427D/5 (1 juvenile), 81 0428D/4 (2), 810501G/ 
4 (1 juvenile), 8105011/1 (1), 810502D/1 (1, broken). 

Collection sites. Muscat, Oman; Unawatuna, nr Galle, Sri 
Lanka; Rubiah, Klah, Ug Murung, Ug Tapa Gadja, Ug Seukundo, 
Pula We, Sumatra. 

Habitat and depth. Subtidal rock, subtidal rock / sand, subtidal 
rock / coral, coral reef; 0-20 m. 

Remarks. A number of these specimens are juveniles, as small as 
10 mm h.d., and with banded spines. The characteristic elongate, 
tridentate pedicellariae of D. setosum (see Clark & Rowe, 1971 ) are 



80 



A.R.G. PRICE AND F.W.E. ROWE 



absent from all specimens examined. The remaining character of the 
red (or in some cases a faded, cream) ring on the anus is the only 
means of distinguishing this species from D. savignyi. In the field 
the two species are easily distinguished by the colour pattern, D. 
savignyi lacking the red ring and having characteristic, irridescent 
blue lines along the upper interamulacra. 

3. Echinothrix calamaris (Pallas, 1774) 

See. Clark & Rowe, 1971: 140; 153; Clark & Courtman Stock, 
1976: 226; Rowe & Gates, 1995: 208. 

Material. 801111A/2 (1), 810422B/1 (1), 810425C/4 (1), 
810426B/4 (1), 810426B/13 (1), 810501K/2 (1), 810501K/3 (1). 

Collection sites. Muscat, Oman; S. Sabang Bay (to S. of Klah), 
E. Sabang Bay, Rubiah, Ug Seukundo, Pula We, Sumatra. 

Habitat and depth. Subtidal rock, subtidal rock / coral / sand, 
coral reef; 0-20 m. 



Family STOMECHINIDAE 

4. Stomopneustes variolaris (Lamarck, 1816) 

SEE. Clark & Rowe, 1971: 140; 153; Clark & Courtman Stock, 
1976: 228; Sloan et al., 1976: 118; Rowe & Gates, 1995: 246. 

Material. 810123B/4 (1). 

Collection sites. Galle, Sri Lanka. 

Habitat and depth. Subtidal rock; 5 m. 

Family TEMNOPLEURIDAE 

5. Mespilia globulus (Linnaeus, 1758) 

See. Clark & Rowe, 1971: 140; 155; Rowe & Gates, 1995: 250. 

Material. 810502F/5 (1). 

Collection sites. Klah, Pula We, Sumatra. 

Habitat and depth. Subtidal rock; 5 m. 

6. Microcyphus ceylanicus Mortensen, 1942 

SEE. Clark & Rowe, 1971: 140; 156. 
Material. 820204A/11 (1). 
Collection sites. SW Kalpitiya, Sri Lanka. 
Habitat and depth. Coral reef; 3-5 m. 

7. Salamacis bicolor L. Agassiz, 1 846 

See. Clark & Rowe, 1971: 140; 156; Clark & Courtman Stock, 
1976: 232. 

Material. 810126B/1 (1). 

Collection sites. Galle, Sri Lanka. 

Habitat and depth. Coral reef; 4-5 m. 



Family TOXOPNEUSTIDAE 

8. Toxopneustes piloleus (Lamarck, 1816) 

See. Clark & Rowe, 1971: 142; 156; Clark & Courtman Stock, 
1976: 234; Rowe & Gates, 1995: 258. 

Material. 801 1 14C/1 (1); 810424A/1 (1). 

Collection sites. Muscat, Oman; W. Klah, Pula We, Sumatra. 

Habitat and depth. Coral reef; 10-12 m. 

Family PARASALENIIDAE 

9. Parasalenia gratiosa A. Agassiz, 1 863 

See. Clark & Rowe, 1971: 142; 157; Rowe & Gates, 1995: 233. 
Material. 810422B/5 (2). 

Collection sites. S. Sabang Bay (to S. of Klah), Sumatra. 
Habitat and depth. Coral conglomerate; 2-8 m. 

Family ECHINOMETRIDAE 

10. Echinometra mathaei (de Blainville, 1825) 

See. Clark & Rowe, 1971: 142; 157; Clark & Courtman Stock, 
1976: 239; Sloan et al., 1979: 1 19; Price, 1983: 76; Rowe & Gates, 
1995:211. 

Material. 801111A/1 (5), 810204A/10 (1). 

Collection sites. Muscat, Oman; SW Kalpitiya, Sri Lanka. 

Habitat and depth. Coral reef, rock / coral / sand; 0-5 m. 

Remarks. Two forms of this species are represented. The speci- 
men from Sri Lanka is relatively large, with h.d. = 48.7 mm. It has 4 
pore-pairs per arc, the spines are uniformly pale blue/green and the 
test, when cleaned, is whitish in colour. The 5 specimens from 
Muscat, Oman are smaller, ranging in size from h.d. = 10-21 mm. 
They have 5 pore-pairs per arc, the spines are dark olive green tipped 
with lilac/brown and the test, when cleaned, is greenish in colour. 
Echinometra mathaei is clearly a complex species which is in need 
of critical investigation to determine whether, as it is currently 
identified, it comprises a single species or more than one closely 
related species (see Mortensen, 1943; Tsuchiya & Nishira, 1984). 

11. Echinostrephus molaris (de Blainville, 1825) 

See. Clark & Rowe, 1971: 142; 157; Clark & Courtman Stock, 
1976: 239-240; Sloan etal., 1979: 1 19; Rowe & Gates, 1995: 212. 

Material. 801210B/6 (1), 8 10423 A/5 (1), 810427B/3 (1), 
810427D/5 (1), 810502G/4 (1) 

Collection sites. Chetlat, Lakshaweep (Laccadive) Islands; Ug 
Bau, Lho Pria Laot, Ug Murung, Klah, Pula We, Sumatra. 

Habitat and depth. Coral reef, subtidal rock, subtidal rock / 
coral; 0^M) m. 



INDIAN OCEAN ECHINODERMS 



81 



DISCUSSION 



Echinoderm collections from the Sindbad Voyage have yielded at 
least 44 species of ophiuroids and 1 1 species of echinoids. Species 
totals for each area sampled, with the corresponding zoogeographic 
subdivision used by Clark & Rowe (1971), are given in Table 2. 
Despite the small size of Pula We (c 20 km x 12 km), 32 ophiuroids 
and 7 echinoids (71% of all species recorded) were encountered at, 
although not necessarily restricted to, this island. This high species 
richness is partly a reflection of the sampling intensity in Pula We, 
but is equally or more an indication of high biodiversity known for 
coral reefs in the SE Asia region (Sheppard, 1987; Wells & Price, 
1992). 

Of the ophiuroids collected, O (A.) purpurea and O. trilineata 
were the most common, occurring in more than 20% of the ophiuroid 

Table 2. Ophiuroid and echinoid species numbers for each area of the 
Indian Ocean sampled during the Sindbad Voyage, (total ophiuroid 
species recorded for all regions = 44 ; total echinoid species recorded 
for all regions =11) 



Sampling area and equivalent 
zoogeographic subdivision 



No. of species recorded 
OPHIUROIDS ECHINOIDS 



Oman (SE Arabia) 


2 


4 


S India (W. India & Pakistan) 


1 





Laccadive (Maldive area) 


7 


1 


Sri Lanka (Sri Lanka area) 


7 


I 


Sumatra (Indonesia / East Indies) 


31 


7 



Of the 44 ophiuroid species collected, ten result in new area 
records (Table 3), as follows: W India (Amphioplus (Lymanella) 
sp.); Sri Lanka (Ophiactis modesta, Ophiarachna robillardi, 
Ophiodyscrita instratus); Maldives area (Cryptopelta granulifera, 
Ophiochaeta hirsuta); and Indonesia / East Indies (Amphiura 
(Amphiura) dejectoides, Amphiura (Amphiura) micra, Amphioplus 
(A.) stenaspis, Ophiogymna pellicula). 

Table 3. New area records and previously known distribution of 
ophiuroid species recorded in the Indian Ocean during the Sindbad 
Voyage 



Species 



New area record Previously known distribution 



Amphiura Indonesia/East 

(Amphiura) Indies 

dejectoides 

Amphiura Indonesia/East 

(Amphiura) micra Indies 

Amphioplus (A. ) Indonesia/East 

stenaspis Indies 

Amphioplus West India 

(Lymanella) sp. (Beypore) 

Ophiactis modesta Sri Lanka 



Ophiogymna 

pellicula 
Cryptopelta 

granulifera 
Ophiarachna 

robillardi 
Ophiochaeta 

hirsuta 

Ophiodyscrita 
instratus 



Indonesia/East 

Indies 
Maldives 

Sri Lanka 

Maldives 

Sri Lanka 



Red Sea; E. Africa (Madagascar) 



N. Australia and possibly E. Africa/ 

Madagascar 
N. Australia and possibly E. Africa/ 

Madagascar 
A. (L.) andreae from Indonesia/ 

East Indies, and A. (L.) laevis from 

Indo-West Pacific 
W. India & Pakistan, and eastwards 

from Bay of Bengal to Hawaiian 

Is. but not Philippines, and 

possibly also E.Africa/Madagascar 
Bay of Bengal and N. Australia 

Mascarene Is. (Mauritius, Reunion, 
Rodrigues group) and N. Australia 
Mascarene Is. 

Is. of W. Indian Ocean, Indonesia/ 
East Indies, N. Australia and S. 
Pacific Is. 

Japan and New Caledonia 
(SW Pacific) 



samples. These species also occupied a wide range of substrata and 
depths (2-30m). Other species occurred in less than 10% of the 
ophiuroid samples, and generally occupied fewer habitats and a 
narrower depth range. The echinoid fauna was less diverse, although 
some species were very common, in particular D. setosum, E. 
calamaris and E. molaris (a coral rock borer), which occurred in 
36%, 19% and 14 % of the echinoid samples respectively. These 
echinoids were found in a wide range of habitats and depths, up to 40 
m in the case of E. molaris. A more comprehensive ecological 
analysis of echinoderms of Pula We, Sumatra is to be undertaken 
following completion of the taxonomic appraisal of the crinoids. 



Acknowledgements. We wish to thank the staff at the Natural History 
Museum, London, for access to the collections and other facilities. We are 
grateful of Dr R. Dalley, P. Hunnam, P. Dobbs and D. Tattle for their 
considerable assistance during field work. One of us (A.R.G.P.) would also 
like to thank T. Severin, leader of the Sindbad Voyage, for the kind invitation 
to participate in the expedition which was made possible by generous support 
from the Ministry of Natural Heritage and Culture, Sultanate of Oman. 
Thanks are due to L. Marsh for identifying several ophiuroids. Financial 
assistance to A.R.G.P. from the Leverhulme Trust is gratefully acknowl- 
edged. 



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Ophiocoma (Echinodermata: Ophiuroidea) from the west coast ofThailand, Andaman 

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Cherbonnier, G. & Guille, A. 1 978. Faune tie Madagascar. 48. Ophiurides. Centre De 

La Recherche Scientifique, Paris: CNRS. 
Clark, A.M. 1 953. A revision of the genus Ophionereis. Proceedings of the Zoological 

Society of London 123(1): 65-94. 
1967. Notes on the family Ophiotrichidae (Ophiuroidea). Annals and Magazine 

of Natural History 9: 637-655. 

1968. Notes on some tropical Indo- Pacific ophiotrichids and ophiodermatids 

(Ophiuroidea). Bulletin of the British Museum (Natural History ), Zoology 16(7): 
275-322. 

Clark, A.M. & Courtman-Stock, J. 1976. Echinoderms of Southern Africa. 277 p. 
London: British Museum (Natural History), London. 

Clark, A.M. & Rowe, F.W.E. 1971. Monograph of Shallow-Water Indo-West Pacific 
Echinoderms. 238 p. British Museum (Natural History), London. 

Clark, H.L. 1938. Echinoderms from Australia. Memoirs of the Museum of Compara- 
tive Zoology. Hansard 55 : 1-596. 

Devaney, D.M. 1970. Studies on ophiocomid brittlestars. 1. A new genus (Clarkcoma) 
of Ophiocominae with evaluation of the genus Ophiocoma. Smithsonian Contribu- 
tions to Zoology No. 51: 1-41. 

1974. Shallow-water asterozoans of southeastern Polynesia 2. Ophiuroidea. 

Micronesica 10: 105-204. 

Guille, A. & Vadon, C. 1985. Les ophiures littorales de Nouvelle-Caledonie. Bulletin 
du Museum National d'Histoire Naturelle, Paris (4)7, Sect. A., no. 1: 61-72. 

Hoggett, A.K. 1991. The genus Macrophiothrix (Ophiuroidea: Ophiotrichidae) in 
Australian waters. Invertebrate Taxonomy 4: 1077-1046. 

1992. Taxonomic and systematic position of the brittlestar genus Macrophiothrix 

H.L. Clark (Echinodermata: Ophiuroidea). PhD Thesis, University of Queensland, 
xviii + 412pp. 

James, D.B. 1989. Echinoderms of Lakshadweep and their zoogeography. Bulletin of 
the Centre for Marine Fisheries Research Institute 43: 97-144. 

Lincoln, R.J. & Shields, J.G. 1979. Invertebrate Animals: Collection and Preserva- 
tion. 150 p. British Museum (Natural History) & Cambridge University Press. 

Marsh L.M. & Price, A.R.G. 1991. Indian Ocean echinoderms collected during the 
Sindbad Voyage (1980-81): 2. Asteroidea. Bulletin of the British Museum (Natural 
History ). Zoology 57(1): 61-70. 



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Mortensen,T. 1 943. A Monogaph of the Echinoidea. III. Echinidae, Strongylocentridae, 

Paraseleniidae. Echinometridae. Reitzel, Copenhagen, pp. 1—446. 
Murakami, S. 1943. Reports on the ophiurans of Palao, Caroline Islands. Journal of 

the Department of Agriculture, KyiisyU Imperial University 7(4): 159-204. 
Murakami, S. 1944. Note on the ophiurans of Amakusa, Kyiisyii. Journal of the 

Department of Agriculture, Kyiisyii Imperial University 7(8): 259-280. 
Nagabhushanam, A.K. & Rao, G.C. 1972. An ecological survey of the marine fauna 

of Minicoy Atoll (Laccadive Archipelago Arabian Sea). Mitteilungen Zoologisches 

Museums Berlin 48(2): 265-324. 
Price, A.R.G. 1983. Echinoderms of Saudi Arabia. Echinoderms of the Arabian Gulf 

coast of Saudi Arabia. Fauna of Saudi Arabia 5 : 28-108. 
Price A.R.G. & Reid, C.E. 1985. Indian Ocean echinoderms collected during the 

Sindbad Voyage (1980-81): 1. Holothurioidea. Bulletin of the British Museum 

(Natural History ), Zoology 48(1): 1-9. 



Rowe, F.W.E. & Gates, J. 1995. Echinodermata. In Wells, A. (ed.) Zoological Cata- 
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Sheppard, C.R.C. 1987. Coral species of the Indian Ocean and adjacent seas: a 
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Sloan N.A., Clark A.M. & Taylor, J.D. 1979. The echinoderms of Aldabra and their 
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Tsuchiya, M. & Nishihara, M. 1984. Ecological distribution of two types of the sea- 
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Vail, L.L. & Rowe, F.W.E. 1989. Status of the genera Ophiopeza and Ophiopsammus 
(Echinodermata: Ophiuroidea) in Australian waters, with the description of a new 
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Bull. mil. Hist. Mas. Land. (Zool.) 62(2): 83-99 



Issued 28 November 1996 



Rare cyclopoid copepods (Crustacea) from 
Mediterranean littoral caves 



D. JAUME AND G.A. BOXSHALL 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK 

SYNOPSIS. Three cyclopoid copepods are reported from anchihaline cave habitats on Mallorca. Both sexes of Cyclopina esilis 
Brian are redescribed. Sexual dimorphism in the mandibular exopod. as discovered in C. esilis, has not previously been reported 
for any cyclopinid. The male of the primitive marine cyclopid Neocyclops (Protoneocyclops) mediterraneus (Kiefer) is described 
in detail for the first time and the characters distinguishing this species are discussed. Both sexes of Euryte longicauda Philippi 
are redescribed. The genus Euryte Philippi is briefly reviewed and the characters used to differentiate species are critically 
reevaluated. It is concluded that all three genera, the cyclopinid genus Cyclopina Claus and the primitive cyclopids Neocyclops 
Gurney and Euryte, are in urgent need of revision, but that this process will be hampered by the inadequacy of published 
descriptions and the lack of available type material. 



INTRODUCTION 



Anchihaline cave habitats are rich sources of interesting and unusual 
crustaceans. Copepods of immense phylogenetic significance have 
been discovered in such caves in the last decade, including the 
platycopioid Antrisocopia Fosshagen, the misophrioid 
Speleophriopsis Jaume & Boxshall, and the calanoid Erebonectes 
Fosshagen (Fosshagen & Iliffe, 1985; Jaume & Boxshall, 1996). 
There have been few reports of cyclopoid copepods in anchihaline 
caves. Recently, however, Rocha & Iliffe (1991, 1994) described a 
new family of cyclopoids, the Speleoithonidae, and the primitive 
cyclopid Troglocyclops, from caves on the Bahamas Islands. During 
our studies of the copepod fauna of Mallorcan caves we discovered 
numerous cyclopoids, including the three rare species described in 
this account. All three species were originally described from the 
Mediterranean Basin. All have been the subject of considerable 
taxonomic confusion because their original descriptions were inad- 
equate and we have attempted to clear up some of this confusion at 
the same time as presenting full redescriptions. 



THE CAVES 



The copepods were collected from four caves located on the East 
coast of Mallorca, less than 20 m inland. These caves occur in two 
different types of substratum. Cova 'C de Cala Varques and Es 
Secret des Moix are in Tortonian (10 Myr BP), coral reef-derived, 
porous calcarenites and mixing-zone corrosion processes seem to 
have played an important role in their development. Cova de na 
Barxa and Cova de na Mitjana are in Triassic, fissured limestones. 
All these caves have subaerial entrances; the difficulty of access can 
be deduced from their topographies, published elsewhere (see be- 
I low). The water conditions varied from cave to cave: In Es Secret des 
Moix, the sampled lake (that located closest to the entrance) was 
completely marine (i.e., in water salinity, a detectable swell, the 
nature of the accompanying fauna), as was the lake in Cova de na 
Mitjana. Cova 'C de Cala Varques and Cova de na Barxa are typical 
anchihaline caves (in the sense of Stock et al., 1986), with a thin 
layer of fresh water on the top of the deeper saline waters of the 
lakes. 



Sampling was undertaken using meat-baited traps placed at dif- 
ferent depths in the cave lakes and left for several days, and by using 
a hand-held plankton net with an extensible handle. The terminology 
used in the descriptions follows Huys & Boxshall (1991). 



SYSTEMATICS 



Order CYCLOPOIDA Burmeister, 1834 
Family CYCLOPINIDAE Sars, 1913 
Subfamily CYCLOPININAE Kiefer, 1927 
Genus Cyclopina Claus, 1863 



Cyclopina esilis Brian, 1938 

(Figs 1-4) 

Cyclopina cf. kieferi: Herbst (1953; 1962) 

Material examined. Cova de na Mitjana (Capdepera). UTM 
coordinates: 539.10; 4390.95. Topography published by Gineset al. 
(1975): 96 individuals, both sexes (BMNH 1995. 1331-1340). 
Collected by D. Jaume, 17 July 1994. 

ADULT female. Body (Figs 1A, B) cyclopiform, up to 0.37 mm 
long. Prosome 5-segmented, about 1.7 times longer than urosome. 
Rostrum developed, oval. Posterolateral margins of cephalosome 
vaulted. First pedigerous somite free, partially concealed by dorsal 
and posterolateral extensions of cephalosome. Second to fourth 
pedigerous somites with evenly rounded posterolateral angles. 
Urosome 5-segmented, with genital and first abdominal somites 
completely fused to form genital double-somite. Serrate hyaline frill 
adorning posterodorsal margin of fifth pedigerous somite, posterior 
margins of genital double-somite and abdominal somites 2 and 3, 
and posterolateral margins of anal somite; degree of serration vary- 
ing directly with body size. Genital double-somite (Fig. ID) 
symmetrical, 1 .6 times longer than wide, expanded anteriorly. Sin- 
gle copulatory pore opening mid-ventrally at about two-fifths of 
distance along double-somite. Paired gonopores located laterally, 
each covered by operculum armed with short spinous process, 1 



©The Natural History Museum, 1996 



84 



D. JAUME AND G.A. BOXSHALL 




Fig. 1. Cyclopina esilis Brian, 1938, adult female. A, body, dorsal view; B, lateral; C, antennule; D, genital double-somite, lateral; E, fifth leg; F-G, dorsal 
view of anal somite and caudal rami, showing variation with body size in the position of the lateral seta of the caudal ramus. 



seta, and 1 long flanged spine (Fig. ID). Anal somite (Figs IF, G) 
bearing smooth operculum. Caudal rami longer than anal somite, 
about 2.6 to 3.3 times longer than wide; proportional length related 
to body size. Armature consisting of 6 setae; position of seta II 



variable, ranging from two-fifths (Fig. 1 G) to three-fifths of distance j 
along distal margin (Fig. IF). 

Antennules (Fig. 1C) symmetrical, 10-segmented, shorter than 
prosome (Fig. IB). Segmental fusion pattern and armature as fol- 



CYCLOPOIDS FROM LITTORAL CAVES 



85 




Fig. 2. Cyclopina esilis Brian, 1938, A and C-F, adult female, B. adult male. A, antenna; B, mandible; C, maxillule; D. maxilla; E, maxilliped; F, 
mandibular exopod. 



lows: segment 1 (corresponding to fused ancestral segments I and 
II), 3 setae; segment 2 (corresponding to fused ancestral segments III 
toV), 5 setae; segment 3 (fused segments VI to IX), 8 setae; segment 
4 (fused segments X and XI), 4 setae; segment 5 (fused segments XII 
to XIV), 6 setae; segment 6 (fused segments XV to XX, partial 
suture present between segments XVI and XVII), 6 setae + aesthetasc; 
segment 7 (fused segments XXI to XXIII), 3 + aesthetasc; segment 
8 (XXIV), 2 setae; segment 9 (XXV), 1 + aesthetasc; segment 10 
(fused segments XXVI to XXVIII), 7 + aesthetasc. 

Antenna (Fig. 2A) 4-segmented. Fused coxa and basis short, 
about 1.5 times longer than wide, armed with 1 inner basal seta 
distally and 1 long seta (representing exopod) on outer margin. 



Endopod 3-segmented. Proximal segment with 1 seta at about two- 
thirds of distance along inner margin; segment covered by long 
spinules on anterior surface. Second segment with 2 lateral and 3 
distal setae (one of them claw-like) along inner margin. Distal 
segment with 7 distal setae, one of them claw-like. Secondary 
ornamentation on segments as figured. 

Mandible (Figs 2B, F) with gnathobase armed with 10 unequal, 
sharp teeth plus 1 dorsal seta; row of 13 spinules located subdistally. 
Palp well developed; basis elongate, with patch of setules and 1 
subdistal seta along inner margin. Exopod (Fig. 2F) inserted at about 
midway of distance along outer margin of basis; 4-segmented; setal 
formula 1,1,1,2; distal, brush-like seta somewhat shorter and thicker 



86 



D. JAUME AND G.A. BOXSHALL 




Fig. 3. Cyclopina esilis Brian, 1938, adult female swimming legs, posterior view. A, leg 1; B, leg 2; C, leg 3; D, leg 4. 



than others, with brush of 6 setules on tip. Endopod 2-segmented, 
shorter than exopod, setal formula 3,6. 

Maxillule (Fig. 2C) with well developed praecoxal arthrite, armed 
with 9 thick, unequal spines around distal margin plus isolated seta 
on posterior surface. Coxa and basis fused: coxal epipodite repre- 
sented by 2 unequal setae; coxal endite represented by small 
cyclindrical knob armed with 1 seta. Proximal and distal endites of 
basis discrete, bearing 3 and 2 setae respectively. Endopod 1- 
segmented, bearing 7 setae. Exopod 1 -segmented, armed with 4 
distal setae and marginal row of long setules. 



Maxilla (Fig. 2D) well developed, 4-segmented. Praecoxa and 
coxa partially fused, endites with setal formula 3,1,3,3. Basis with 
large endite bearing claw-like spine plus 2 unequal setae. Endopod 
2-segmented; proximal segment representing fused first and second 
ancestral endopodal segments; distal representing fused third and 
fourth ancestral segments; endopod setation formula (2+2),(2+4). 

Maxilliped (Fig. 2E) slender, 6-segmented. Praecoxa and coxa 
fused forming syncoxa, bearing 3 (coxal) endites with setal formula 
1,3,2. Basis with medial margin swollen, ornamented with marginal 
row of long setules; 2 setae implanted subdistally on medial margin. : 



CYCLOPOIDS FROM LITTORAL CAVES 



87 




Fig. 4. Cyclopina esilis Brian, 1938, adult male. A, body, dorsal; B, lateral; C, genital somite, ventral; D, antennule; E, fifth leg. 



Endopod 4-segmented, setal formula 0,0,1,4; transverse row of 
setules on segment 3. 

Swimming legs 1 to 4 (Fig. 3) biramous, both rami 3-seg- 
mented. Legs subequal in size except first, somewhat reduced. 
All legs richly ornamented with denticles, as figured; anterior 
surface of coxae covered by small denticles, omitted from fig- 
ures. Spines on exopodal segments flanged with serrate hyaline 
frill; distal spine on third exopodal segment of legs 1 and 2, and 
spines on endopod of leg 4 flanged only on one side. Armature as 
follows: 



Coxa 



Basis Exopod 



Endopod 



Legl 


0-1 


1-1 


I-1;I-1;III,I,4 


0-l;0-l; 1,2,3 


Leg 2 


0-1 


1-0 


I-1;I-1;III,I,5 


0-1 ;0-2; 1,2,3 


Leg 3 


0-1 


1-0 


I-1;1-1;III,I,5 


0-1 ;0-2; 1,2,3 


Leg 4 


0-1 


1-0 


I-1;I-1;II,I,5 


0-1 ;0-II; 1,2,1+1 



Fifth legs (Fig. IE) uniramous, 2-segmented, joined by naked 
intercoxal sclerite. Coxa and basis fused forming trapezoidal 
protopodal segment; inner margin with row of long setules; outer 
margin with long, smooth seta subdistally. Distal segment (exopod) 
about 1.6 times as long as wide, produced distally into median 



D. JAUME AND G.A. BOXSHALL 



process bearing 1 long, plumose seta; 1 flanged spine present 
subdistally on each side of process, outer stouter, longer than 
segment; inner spine less than half length of outer spine; spinule 
ornamentation on segment as figured. 

Adult male. Body (Figs 4A, B) up to 0.38 mm long, more 
slender than female. Urosome 6-segmented, with genital somite 
(Fig. 4C) symmetrical, slightly expanded laterally; paired gonopores 
opening ventrally at posterior border of somite; genital opercular 
flaps each armed with tiny inner spine plus 2 long, outer setae. 

Antennules (Fig. 4D) 15-segmented, symmetrical, digeniculate. 
Geniculations between segments homologous with ancestral seg- 
ments XV and XVI (9 and 10), and between XX and XXI (13 and 
14). Segment 9 (XV) cup-shaped, forming sheath around proximal 
half of segment 10 (XVI). Segmental fusion pattern and armature as 
follows: segment 1 (corresponding to fused ancestral segments I and 
II), 3 setae + aesthetasc; segment 2 (fused ancestral segments III to 
V), 5 setae; segment 3 (fused ancestral segmentsVI andVII), 4 setae; 
segment 4 (VIII), 2 setae; segment 5 (partially fused ancestral 
segments IX to XI), 6 setae; segments 6 to 9 (XII to XV), 2 setae 
each; segment 10 (XVI), 1 pectinate spine, 1 seta + aesthetasc; 
segments 1 1 and 12 (XVII and XVIII), 1 pectinate spine and 1 seta 
each; segment 13 (fused ancestral segments XIX and XX), 1 pecti- 
nate spine, 1 modified flattened spine, and 1 seta; segment 14 (fused 
ancestral segments XXI and XXII), 1 modified flattened, spine 
plate, 1 seta + aesthetasc; segment 15 (fused ancestral segments 
XXIII to XXVIII), 1 1 + aesthetasc. 

Segmentation and setation of other cephalosomic appendages and 
swimming legs 1 to 4 as in female, except mandibular palp (Fig. 2B); 
distal brush-like seta on exopod much shorter and thicker, setules on 
tip longer than in female. 

Fifth legs (Fig. 4E) resembling female condition, but with 2 
additional setae implanted subdistally along inner margin of exopod. 

Remarks. The Cyclopina from the cave on Mallorca belongs to 
the group of species in the genus that displays a female leg 5 with the 
inner spine of distal segment less than half the length of the outer 
spine, the latter being longer than the segment itself. This group 
comprises Cyclopina esilis Brian, 1938, C. americana Herbst, 1982, 
and C. cuipora Lotufo, 1994. The taxon from Mallorca differs 
clearly from C cuipora. The female antennule is 10-segmented (not 
12-segmented as in C. cuipora), and the intercoxal sclerite of leg 4 
is almost completely smooth (not powerfully ornamented with 
several rows of thick spinules as in C. cuipora) (Lotufo, 1994). 
Differences from C americana include the short, subquadrate cau- 
dal rami (Herbst, 1982) which contrast with the elongate (2.5 to 3.2 
times as long as wide) caudal rami of the Mallorcan taxon. 

The Cyclopina from Mallorca is identified as C. esilis, based on 
the segmentation of the female antennule, the setation of leg 5 and 
the proportions of the caudal rami. We noted variation in the 
proportional length of the caudal rami within the Mallorcan popula- 
tion. A similar degree of variability in length of the caudal rami has 
been reported in C. esilis (Brian, 1938; Monchenko, 1979). 

Apparent differences in the armature of mouthparts have not been 
evaluated since we suspect that the armature of mouthparts in C. 
esilis (as in most species of Cyclopina) were inadequately described 
in the original descriptions. This is evident in the presence of a coxal 
endite, armed with 1 seta, on the maxillule of the Cyclopina from 
Mallorca. This character state (presence of coxal endite) had never 
previously been noted in Cyclopina and was known only from 
Cyclopinodes elegans (T Scott, 1894) in the family Cyclopinidae 
(Huys & Boxshall, 1991 ). The coxal endite may well be present on 
the maxillule of all Cyclopina species. It is present in C. gracilis 
Claus, 1863 from the coast of Scotland (BMNH 1986.377) (pers. 



obs.), and perhaps also in C oblivia Monchenko, 198 1 , according to 
Monchenko (1989: Fig. 9). 

The Cyclopina reported by Herbst (1953; 1962) from Banyuls 
(South France) and from Brittany (NW France) as C. cf. kieferi 
Schafer, 1936 was recorded living as a commensal with polychaetes 
(Bretagne) and free-living in the marine interstitial (Banyuls). The 
illustrations provided by Herbst differ from the original description 
by Schafer (1936) in an important character for the taxonomy of the 
genus, the relative length of the female leg 5 exopodal spines. This 
discrepancy was already noted by Lotufo (1994) when he presented 
the differential diagnosis of C. yutimaete Lotufo in comparison to C. 
kieferi. As the female leg 5 of C. cf. kieferi figured by Herbst (1953; 
1962) is identical to that of C. esilis, we here consider Herbst's 
material as belonging to C. esilis. 

The distribution of Cyclopina esilis thus encompasses the littoral 
zone from the Black Sea to the western approaches of the English 
Channel; a distribution equivalent to the Mediterranean and 
Lusitanian provinces of classical marine biogeography (Ekman, 
1953). 



Family CYCLOPIDAE Dana, 1846 
Subfamily HALICYCLOPINAE Kiefer, 1927 
Genus Neocyclops Gurney, 1 927 



Neocy clops (Protoneocyclops) mediterraneus (Kiefer, 
1960) 

(Figs 5-7) 

Pareuryte mediterranea: Kiefer (1960). 

Neocyclops remanei mediterraneus: Pesce & Galassi (1987) 

Material examined. Cova 'C' de CalaVarques (Manacor). UTM 
coordinates: 525.27; 4372.19. Topography in Trias & Mir (1977): 
Two adult males (one not preserved) and 2 copepodids (one not 
preserved) (BMNH 1995. 1329-1330). Collected by authors, 29 
March 1995. 

Adult male. Body (Fig. 5A) cyclopiform, up to 0.58 mm long, 
colourless. Nauplius eye absent. Prosome about 1 .4 times as long as 
urosome, 5-segmented, first pedigerous somite completely con- 
cealed by carapace-like, posterior extension of cephalosome. Rostrum 
(Fig. 5B) triangular in frontal view. Urosome 6-segmented, robust. 
Fifth pedigerous somite with pointed posterolateral angles; entire 
hyaline frill adorning posterodorsal margin of somite. Genital somite 
(Fig. 5C) slightly expanded laterally, with ventrolateral fold each 
side of somite at about one-third distance from posterior margin; 
folds slightly projecting dorsally. Paired gonopores opening ventrally, 
each covered by opercular flap derived from sixth leg; flaps each 
armed with 1 inner flanged spine and 2 outer plumose setae. 
Abdominal somites 1 to 3 subequal, with posterior margins adorned 
with entire hyaline frill. Anal somite bearing operculum dorsally at 
about midlength; operculum ornamented with serrate hyaline frill; 4 
rows of transverse setules adorning sides of anal cleft; posterolateral 
margins of somite bearing serrate hyaline frill. Caudal rami 2.6 
times as long as wide, inserted widely separate from each other; 
secondary ornamentation of pores and tiny spinules distributed as 
figured; armature consisting of 7 setae; seta I reduced, tiny, im- 
planted ventrolaterally about one-third of distance along ramus; seta 
II implanted dorsolateral^ at about three-quarters of distance along 
ramus. 

Antennules (Fig. 5E) 1 6-segmented, not extending beyond poste- 
rior margin of prosome, symmetrical, digeniculate (Fig. 5A). 



CYCLOPOIDS FROM LITTORAL CAVES 



J" 0.05 mm 
QD 

-£_ P 0.1 25 mm 




Fig. 5. Neocyclops (Protoneocyclops) mediterraneus (Kiefer, 1960), adult male. A, body, dorsal; B, detail of rostral plate; C, genital somite, ventral; D, 
anal somite and caudal rami, dorsal; E, antennule; F, fifth leg. 



Geniculations between segments homologous with ancestral seg- 
ments XV and XVI (10 and 1 1), and between XX and XXI (14 and 
15). Segment 10 cup-shaped, forming sheath around proximal half 
of segment 1 1 . Segmental fusion pattern and armature as follows: 
segment 1 (corresponding to fused ancestral segments I to V), 8 setae 
+ 3 aesthetascs; segment 2 (partially fused ancestral segments VI and 
VII), 4 setae; segment 3 (VIII), 2 setae; segment 4 (IX), 2 setae + 



aesthetasc; segments 5 to 8 (X to XIII), 2 setae each; segment 9 
(XIV), 2 setae + aesthetasc; segments 10 and 11 (XV and XVI), 2 
setae each; segment 12 (XVII), 2 setose spines; segment 13 (XVIII), 
1 setose spine, 1 seta + aesthetasc; segment 14 (fused segments XIX 
and XX), 1 modified flattened spine and 3 setae; segment 15 
(partially fused segments XXI and XXII), 2 modified flattened 
spines, 1 seta + aesthetasc; segment 16 (fused segments XXIII to 



90 



D. JAUME AND G.A. BOXSHALL 




Fig. 6. Neocyclops (Protoneocyclops) mediterraneus (Kiefer, 
maxilliped. 



0.05 mm 



I960), adult male. A, antenna; B, labrum. ventral; C, mandible; D. maxillule; E, maxilla; F. 



XXVIII), 11+2 aesthetascs. 

Antenna (Fig. 6A) well developed, 5-segmented. Coxa and basis 
separate, coxa small, unarmed. Basis with 2 distal setae on inner 
margin and long distal seta, representing exopod, on outer margin. 
First endopod segment with seta inserted midway along inner mar- 
gin. Second endopod segment with 2 lateral and 3 distal setae along 
inner margin; outer margin covered by patch of setules. Third 
endopod segment with 7 unequal setae on tip; outer margin adorned 
with 2 patches of setules, as figured. 

Labrum (Fig. 6B) with laterally serrate distal margin and row of 9 
rounded teeth midway along margin; paired transverse rows of long 



spinules located subdistally on anterior surface of labrum. 

Mandible (Fig. 6C) with coxal gnathobase armed with 9 sharp, 
unequal teeth, 2 dorsal setae, and 1 naked setiform element located 
between third and fourth ventralmost teeth; inner dorsal seta with 
long spinules along one side; outer dorsal seta longer, with short 
spinules on both sides; transverse row of 7 long spinules disposed 
subdistally on cutting blade. Mandibular palp reduced to knob 
bearing 3 unequal setae. 

Maxillule (Fig. 6D) with well developed praecoxal arthrite; arma- 
ture consisting of 3 stout spines, 1 tiny spine and 3 slender setae 
proximally, and distal lobe armed with 4 thick, denticulate spines; 2 



CYCLOPOIDS FROM LITTORAL CAVES 



91 




Fig. 7. Neocyclops (Protoneocyclops) mediterraneus (Kiefer, 1960), adult male swimming legs, posterior view. A, leg 1; B, leg 2; C, leg 3; D, leg 4. 



patches of tiny denticles on segment, as figured. Palp comprising 
coxobasis with 1 spinulate spine and 2 setae distally and 1 seta 
(representing exopod) implanted on outer margin, and 1 -segmented 
endopod bearing 3 setae (obscured in Fig. 6D). 

Maxilla (Fig. 6E) 4-segmented, powerfully developed. Praecoxa 
and coxa separate. Praecoxa with single, distal endite armed with 2 
elements. Proximal coxal endite represented by single seta; distal 
coxal endite discrete, armed with 1 spine fused to endite and 
ornamented with 2 strong spinules, and 1 pectinate spine. Basis with 
endite bearing 2 stout pectinate elements, one fused to segment, plus 



reduced pinnate seta. Endopod 1 -segmented, bearing 3 stout pecti- 
nate elements plus 2 reduced, unequal setae. 

Maxilliped (Fig. 6F) somewhat reduced in size, 4-segmented. 
Synooxa bearing 2 weakly developed endites with spine formula 
2, 1 . Basis with 2 distal setae on inner margin; secondary ornamen- 
tation of spinules and setules on segment as figured. Endopod 
2-segmented, setal formula 2,3. Ornamentation on spines and setae 
of maxilliped as figured; setae on distal endopodal segment naked. 

Swimming legs 1 to 4 (Fig. 7) biramous, both rami 3-segmented, 
except leg 4 with 2-segmented endopod; distal endopodal segment 



92 



D. JAUME AND G.A. BOXSHALL 



subdivided by posterior surface suture marking plane of fusion 
between second and third segments. Legs subequal in size except 
first somewhat reduced. Intercoxal sclerites lacking ornamentation. 
Legs richly ornamented with spinules, setules and denticles, as 
figured. Armature as follows: 



Coxa 



Basis Exopod 



Endopod 



Legl 


0-1 


1-1 


I-1;I-1;III,I,4 


0-l;0-2;I,I+l,3 


Leg 2 


0-1 


1-0 


I-1;I-1;III,I,5 


0-l;0-2;I,II,3 


Leg 3 


0-1 


1-0 


I-1;I-1;III,I,5 


0-l;0-2;I,II,3 


Leg 4 


0-1 


1-0 


I-1;I-1;III,I,IV 


0-1:1,11,111+1 



Spines on legs 1 and 2 adorned with spinules, those on leg 3 
flanged with serrate hyaline frill. Outer spines on exopod and outer 
and distal spines on endopod of leg 4 ornamented with serrate 
hyaline frill; inner margin setae on both rami modified, spine-like, 
ornamented with short pinnules proximally and serrate membrane 
distally. 

Fifth legs (Fig. 5F) 4-segmented, joined by smooth intercoxal 
sclerite. Coxa and basis separate, coxa unarmed, basis bearing 
plumose seta on outer margin. First exopodal segment elongate, 
about as long as coxa and basis combined, outer margin with 
subdistal spine flanged with serrated hyaline frill; spinous process 
just anterior to insertion point of spine; flanged spine longer than 
segment; inner margin of segment bearing 1 distal spine adorned 
with sparse setules and about as long as segment. Distal margin of 
second exopodal segment bearing 1 seta flanked by 2 spines flanged 
with serrate hyaline frill; spinous process just proximal to insertion 
point of outer spine; outer spine slightly longer than inner, and 
longer than first exopod segment; inner spine about as long as first 
endopod segment; seta shorter than spines. Secondary ornamenta- 
tion on fifth leg segments as figured. 

Remarks. The genus Neocyclops Gurney, 1927 contains 15 spe- 
cies distributed in coastal waters of the Northeast and Tropical 
Atlantic (including the Caribbean), the Mediterranean, the Black 
and Red Seas, the Indian Ocean, as well as the Pacific (Papua New 
Guinea) (Petkovski, 1986; Fiers, 1986; Pesce & Galassi, 1993; 
Lotufo & Rocha, 1993; Rocha, 1995). Petkovski (1986) has split the 
genus into two subgenera according to the number of exopodal 
segments of the male fifth legs. The subgenus Neocyclops, charac- 
terized by a 3-segmented male leg 5, embraces the following species: 
N. medius Herbst, 1955, A', vicinus (Herbst, 1955), N. affinis (Plesa, 
1961), N. salinarum (Gurney, 1927) and N. remanei (Herbst, 1952). 

The subgenus Protoneocyclops, with 4-segmented male fifth legs, 
comprises P. stocki Pesce, 1985, P. geltrudeae Pesce & Galassi, 
1993, P. papuensis Fiers, 1986, P. mediterraneus (Kiefer, 1960), P. 
herbsti Petkovski, 1986, P. wellsi Petkovski, 1986 and P. ferrarii 
Rocha, 1995. This subgenus displays the so-called full Tethyan 
pattern of distribution (Stock, 1993), i.e., circum-tropical in the 
entire region of the former Tethys Sea. 

Three other species, viz. N. impwvisus Plesa, 1973 from Cuba, 
and N. magnus (Sewell, 1949) and N. parvus (Sewell, 1949) from 
islands in the Indian Ocean, cannot be assigned to either subgenus as 
their males are unknown. 

Three representatives of the genus are known so far from the 
Mediterranean region. Neocyclops (N.) salinarum, originally de- 
scribed from the Suez Canal, was reported also from the Camargue 
(South France) and the Sirbonian lagoon (Mediteranean coast of 
Sinai) (Gurney, 1927a; 1927b;Aguesse&Dussart, 1956; Por, 1973). 
As Petkovski (1986) pointed out, the identity of the French popula- 
tion needs to be confirmed. Similarly the single copepodid from the 



Andaman Islands (Indian Ocean), assigned by Sewell (1949) to this 
species, should be reexamined. Neocyclops (A 7 .) vicinus, a species 
distributed along the coasts of Brazil and the Lesser Antilles (Pesce 
& Galassi, 1993; Lotufo & Rocha, 1993), has been also reported 
from the Black Sea (as Eurycyclops remanei vicinus) by Plesa 
(1963) and Monchenko (1975). As pointed out by Lotufo & Rocha 
(1993), this record is dubious since their material seems more 
closely related to N. (N.) remanei than to N. (A 7 ,) vicinus. 

The single representative of the subgenus Protoneocyclops in 
Mediterranean waters is Neocyclops (P.) mediterraneus, originally 
described by Kiefer (1960) as Pareuryte mediterranea from an 
anchihaline cave on Menorca (Balearic Islands). Later, Pesce & 
Galassi (1987) reported it from an anchihaline cave in Southern 
Italy. Plesa ( 1981 ) cited the same species from Cuba, but this record 
has been reassigned by Petkovski (1986) to N. (N.) stocki, a taxon 
widespread in the Caribbean region (Pesce & Galassi, 1993). 

The Neocyclops from Mallorca has been identified on the basis of 
the 4-segmented condition of the male fifth legs and the relative 
lengths of the armature elements on this leg. Mallorca is also close 
to the type-locality of the species (Menorca). Other characters could 
not be checked against Kiefer's (1960) original description since this 
contained only 5 drawings (viz. female anal somite and caudal rami, 
distal segment of endopod of female leg 4; fifth leg of both sexes, and 
genital operculum of male). In addition, Kiefer did not designate 
types for the species. Pesce & Galassi (1987) had only 2 females at 
their disposal for their supplementary description. 

A differential diagnosis of Neocyclops (Protoneocyclops) 
mediterraneus (Kiefer, 1960) can be constructed based on charac- 
ters of the male fifth leg. It differs from N. (P.) geltrudeae Pesce & 
Galassi (1993) from Curacao (Antilles) in the number of armature 
elements on the distal segment (3, compared to 4 inA 7 . (P.) geltrudeae). 
Differences from N. (P.) papuensis Fiers, 1986 from New Guinea 
and N. (P.) ferrarii Rocha, 1995 from Brazil involve the relative 
lengths of the spines on the distal segment (the inner spine is clearly 
longer than the outer in both these species whereas in N. (P.) 
mediterraneus the outer spine is subsimilar, slightly longer than the 
inner). Differences from N. (P.) herbsti Petkovski (1986) from the 
Red Sea, and N. (P.) stocki Pesce, 1985 from the Caribbean, are 
based on the relative lengths of the flanged spines on the 2 distal 
segments of leg 5 (these are clearly shorter than the first exopodal 
segment whereas in N. (P.) mediterraneus they are longer than the 
segment). In addition, in N. (P.) herbsti the armature element on the 
inner margin of the first exopodal segment is a plumose seta, 
whereas inA 7 . (P. ) mediterraneus it is a thick spine. Differences from 
N. (P.) wellsi Petkovski (1986) from Mozambique lie only in the 
nature of the armature element on the inner margin of the first 
exopodal segment, which is also a seta in this species instead of a 
thick spine. 



Subfamily EURYTEINAE Monchenko, 1975 
Genus Euryte Philippi, 1843 



Euryte longicauda Philippi, 1843 emend. Giesbrecht, 
1900 

(Figs 8-11) 

Material examined. Cova de na Barxa (Capdepera). UTM coor- 
dinates: 539.30; 4393. 10. Topography inAndrewsef al. (1989): Two 
adult females, 1 adult male, and 1 copepodid (BMNH 1995. 1323— 
1326). Collected by authors, 3 April 1995. - Cova de na Mitjana 
(Capdepera): 19 adult males, 7 adult females, and 5 



i 



CYCLOPOIDS FROM LITTORAL CAVES 



93 



0.125mm 




Fig. 8. Euryte longicauda Philippi, 1843, adult male. A, body, dorsal; B, lateral; C, detail of rostral plate; D, genital and first abdominal somites, ventral; 
E, same, lateral; F, anal somite and caudal rami, dorsal; G, antennule; H, fifth leg. 



copepodids (BMNH 1995. 1313-1322). Collected by authors, 1 (BMNH 1995. 1327). Collected by D. Jaume, 25 May 1994. 
April 1995. - Es Secret des Moix (Manacor). Coordinates: 523.69; Adultmale. Body (Figs 8A, B) cyclopiform, up to 0.61 mmlong, 
4365.53. Topography in Gines et al. (1975): One adult female body surface completely covered by tiny cuticular granulations. 



94 



D. JAUME AND G.A. BOXSHALL 




0.05 mm 



Fig. 9. Euryte longicauda Philippi, 1843, adult male. A, antenna; B, mandible; C. maxillule praecoxal arthrite; D. maxillulary palp; E. maxilla; F, 
maxilliped. 



Prosome about 1.4 times longer than urosome, comprising 
cephalosome plus 4 free pedigerous somites; first pedigerous 
somite completely concealed beneath posterior extension of 
cephalosome, with lateral margins weakly sclerotized; second to 
fourth somites with evenly rounded posterolateral corners. Ros- 
trum (Fig. 8C) fused at base, well developed, subtriangular in 
frontal view. Urosome 6-segmented; first abdominal somite with 
pointed posterolateral angles; entire hyaline frill adorning 
posterodorsal margin. Genital somite (Figs 8D,E) symmetrical, 
laterally expanded, with entire hyaline frill around posterodorsal 
margin; pair of gonopores opening ventrally at posterior border of 
somite; opercular flaps each armed with 1 inner flanged spine plus 
2 outer setae. Third to fifth urosome somites subequal, narrower 
than genital somite, with entire hyaline frill adorning posterior 
margin. Anal somite (Fig. 8F) about same size as preceding 
somites; smooth anal operculum present at one third of distance 
along somite; serrate hyaline frill around posterolateral margins of 



somite. Caudal rami (Fig. 8F) of variable length, from 4.4 to 6.4 
times longer than wide, slightly divergent; distal part slightly 
wider; armature consisting of 6 setae; seta II located subdistally. 

Antennules (Fig. 8G) 1 6-segmented, symmetrical, digeniculate 
with geniculations between segments homologous with ancestral 
segments XV and XVI (10 and 11), and segments XX and XXI (14 
and 15). Segment 10 cup-shaped, forming sheath around proximal 
half of segment 1 1 . Segmental fusion pattern and armature as 
follows: Segment 1 (corresponding to fused ancestral segments I to 
V), 8 setae + 3 aesthetascs; segment 2 (corresponding to fused 
ancestral segments VI and VII), 4 setae; segment 3 (VIII), 2 setae; 
segment 4 (IX), 2 + aesthetasc; segments 5 to 8 (X to XIII), 2 setae 
each; segment 9 (XIV), 2 + aesthetasc; segment 10 (XV), 2 setae; 
segment 11 (XVI), 2 + aesthetasc; segment 12 (XVII). 1 short 
denticulate spine and 1 seta; segment 13 (XVIII), 1 short denticulate 
spine, 1 seta + aesthetasc; segment 14 (fused ancestral segments 
XIX and XX), 1 short denticulate spine, 1 modified flattened spine, 



CYCLOPOIDS FROM LITTORAL CAVES 



95 




Fig. 10. Euryte longicauda Philippi, 1843. adult male swimming legs, posterior view. A, leg 1; B. leg 2; C, leg 3; D, leg 4. 



1 seta + minute aesthetasc; segment 15 (fused ancestral segments 
XXI to XXIII), 2 modified flattened spines, 1 seta + aesthetasc; 
segment 16 (fused segments XXIV to XXVIII), 11 setae + 2 
aesthetascs. 

Antenna (Fig. 9A) 4-segmented. First segment representing par- 
tially fused coxa and basis, armed with 2 inner basal setae distally, 
and 1 outer seta representing exopod; patch of setules present 
midway along outer margin of segment. Endopod 3-segmented; first 
segment about as long as coxa and basis combined, armed with 1 
seta at three-quarters of distance along inner margin; outer margin 



ornamented with setules. Segments 2 and 3 subsimilar in size, each 
about half length of segment 1 . Segment 2 armed with 2 lateral and 
3 distal setae, one of latter (seta VIII in scheme of Boxshall & 
Evstigneeva, 1994) claw-like, along inner margin. Segment 3 armed 
with 7 distal setae, one claw-like; outer margin with 2 rows of 
setules, as figured. 

Mandible (Fig. 9B) with large coxal gnathobase bearing 11 
unequal, sharp blades, plus 2 dorsal spines; outer dorsal spine 
spinulate; transverse row of 6 thin spinules located adjacent to 
cutting edge. Palp reduced to knob bearing 3 setae. 



96 



D. JAUME AND G.A. BOXSHALL 



Maxillule with praecoxal arthrite (Fig. 9C) well developed, armed 
distally with 4 stout, denticulate spines, plus 6 more proximal 
elements, ranging from a tiny seta to a thick denticulate spine. Palp 
(Fig. 9D) comprising coxobasis with medial gnathobase-like struc- 
ture and minute endopodal segment bearing 2 setae; distal margin of 
coxobasal gnathobase provided with 9 irregular blades; coxobasis 
with 2 setae located subapically on dorsal margin and single seta, 
representing exopod, located proximally on distal surface. 

Maxilla (Fig. 9E) well developed, 4-segmented. Praecoxa and 
coxa incompletely separate. Praecoxa naked, lacking endites. Coxa 
with proximal endite represented by single seta; distal endite power- 
fully developed, bearing 2 stout, spinulate spines, proximal spine 
bearing single, conspicuous strong spinule on outer margin and row 
of thinner spinules on inner margin. Basis with endite bearing 3 
unequal, claw-like setae. Endopod 1 -segmented, armed with total of 
3 stout spine-like setae, 1 naked seta and 1 very reduced seta. 

Maxilliped (Fig. 9F) 5-segmented, prehensile. Syncoxa bearing 2 
weakly developed endites provided with single seta each. Basis with 
inner margin covered by patch of setules and single seta positioned 
distally. Endopod 3-segmented, first segment short, unarmed; sec- 
ond segment elongate, inner margin covered by patch of setules, 
armed with 2 setae laterally; small distal segment with 2 stout, 
curved claws plus 2 accessory setae. 

Swimming legs 1 to 4 (Fig. 1 0) biramous, both rami 3-segmented. 
Legs subequal in size except first somewhat smaller. Intercoxal 
sclerites lacking ornamentation and getting progressively narrower 
from legs 1 to 4. All spines on segments flanged bilaterally with 
serrate hyaline frill except distalmost spine on exopod of leg 1, 
which is flanged with frill on outer side only, inner side adorned with 
row of setules. Secondary ornamentation and pore pattern on seg- 
ments as figured; pores on coxa and basis possibly overlooked. 
Armature as follows: 



Coxa 



Basis Exopod 



Endopod 



Legl 


0-1 


1-1 


I-1;I-1;IIU,4 


0-l;0-2;I-II-III 


Leg 2 


0-1 


1-0 


I-1;I-1;III,I,IV+1 


0-1;0-2;I-II-III 


Leg 3 


0-1 


1-0 


1-1:1-1 ;III,I,V 


0-1:0-2:1-11-111 


Leg 4 


0-1 


1-0 


1-1:1-1 ;II,I,V 


0-l;0-II;I-II-II 



Fifth legs (Fig. 8H) uniramous, 3-segmented, joined by smooth 
intercoxal sclerite. Coxa and basis separate, former naked, latter 
with single seta on outer margin. Distal segment (exopod) elongate, 
about 2.5 times as long as wide; armature consisting of flanged spine 
as long as segment located two-thirds of distance along outer 
margin, plus 2 flanged spines and single seta on distal margin; distal 
seta shorter than both spines and segment itself; distal spines located 
either side of seta, flanged, both clearly longer than segment, inner 
longer, about 1 .5 times as long as segment. Secondary ornamenta- 
tion of spinules present on outer margin of exopod of some 
individuals, similar to that figured on female leg 5 (Fig. 1 IE), not 
present in figured specimen (Fig. 8H). 

Adult female. Body (Figs 1 1A, B) up to 0.74 mm long, resem- 
bling male. Urosome 5-segmented; genital and first abdominal 
somites partially fused to form genital double-somite. Genital dou- 
ble-somite (Fig. 1 1C) symmetrical, subdivided dorsally by partial 
suture line; single copulatory pore opening mid-ventrally about 
one-third of distance along somite, connected via copulatory duct 
to fused seminal receptacles. Paired gonopores located laterally; 
gonopores covered by opercula, each consisting of lobe projecting 
dorsally bearing 2 setae and 1 tiny spine. Tapering soft lobe point- 
ing posteriorly located at both sides of somite just behind 



operculum. Entire hyaline frill present along posterior margin of 
somite. 

Antennules (Fig. 1 ID) 2 1 -segmented, not extending beyond pos- 
terior margin of cephalosome (Figs 1 1 A, B), symmetrical. Segmental 
fusion pattern and armature as follows: Segment 1 (corresponding to 
fused ancestral segments I to V), 8 setae; segment 2 (fused ancestral 
segments VI and VII), 4 setae; segments 3 to 9 (VIII to XIV), 2 setae 
each; segments 10 to 13 (XV to XVIII), 1 seta each; segment 14 
(XIX), naked; segment 15 (XX), 1 seta; segment 16 (XXI), 1 seta + 
aesthetasc; segment 17 (XXII), naked; segment 18 (XXIII), 1 seta; 
segment 19 (XXIV), 2 setae; segment 20 (XXV), 2 + aesthetasc; 
segment 21 (fused XXVI to XXVIII), 7 + aesthetasc. 

Segmentation and setation of other cephalosomic appendages and 
swimming legs 1 to 4 as in male. 

Fifth legs (Fig. 1 IE) resembling those of male, but with shorter 
exopod, about twice as long as wide; inner distal spine almost twice 
as long as segment; spine on outer margin clearly longer than 
segment. Secondary ornamentation of spinules on outer margin of 
exopod not discernible in some individuals. 

Remarks. The genus Euryte typically contains shallow water 
hyperbenthic species, although Brady (1910) reported the genus 
from depths of 320 m in the Antarctic and some species have been 
found living in the interstices of coarse sand, or in association with 
seaweed or corals. Ten species are currently recognized, distributed 
worldwide (Gurney, 1927b; Sewell, 1949; Herbst, 1989; Humes, 
1991; 1992), with the possible exception of the Pacific coast of 
South America. Apart from the original contributions by Giesbrecht 
(1900) and Sars (1913-1918), new species have been described 
mainly on the basis of a biometric analysis of characters that have 
otherwise proved to exhibit high intra-populational variability (such 
as the relative legth of caudal rami), or that may vary significantly in 
their measurements simply according to the precise angle of obser- 
vation (such as the relative length of the armature elements on the 
distal segment of the endopod of leg 4). Such characters are widely 
used in the two identification keys available for the genus ( Vervoort, 
1964; Herbst, 1989), and their use has resulted in the false impres- 
sion of cosmopolitanism of some taxa (viz. E. longicauda and E. 
robusta Giesbrecht, 1900; see Kiefer (1929) and Sewell (1949)). 

The type material for most species of Euryte is no longer extant. 
This hampers the necessary revision of the genus, that could permit 
the critical reevaluation of all those taxa established on the basis of 
variable characters. 

Using material from the type locality of both species (the Gulf of 
Naples), Giesbrecht (1900) differentiated E. longicauda Philippi, 
1843 fromE. robusta Giesbrecht, 1900 mainly by the proportions of 
the caudal rami and by details of the armature of the male antennule. 
The proportions of the caudal rami of the Mallorcan population 
overlap the characteristic values for both species given by Giesbrecht 
(1900). The armature of the male antennule, however, corresponds 
to that off. longicauda: the cup-shaped segment 10 carries 2 slender 
setae, whereas in E. robusta it carries a characteristic robust, S- 
shaped spine plus a seta. On this basis we have assigned the Euryte 
from the Mallorcan caves to E. longicauda. 

The differential diagnosis separating E. longicauda from Mallorca 
from E. robusta can be completed as follows (see the detailed 
illustrations of the latter species in Huys & Boxshall, 1991): in E. 
longicauda the proximal spine on the distal coxal endite of the 
maxilla of both sexes is armed with a single, strong spinule on one 
side and a row of thinner spinules on the other side; in E. robusta 
both sides are armed with thin spinules. Additionally, in£. longicauda 
there is a transverse dorsal suture midway along the female genital 
double-somite that seems to be absent in E. robusta. 



CYCLOPOIDS FROM LITTORAL CAVES 



97 




Fig. 11. Euryte longicauda Philippi, 1843, adult female. A, body, dorsal; B, lateral; C, genital double-somite, lateral; D, antennule; E, fifth leg. 



The distribution of E. longicauda apparently encompasses all 
European coasts from the Black Sea to the Arctic Ocean (Franz- 
Joseph Land) and the East coast of Greenland (Giesbrecht, 1900; 
Herbst, 1989) but many of the records of this species are accompa- 
nied by inadequate descriptions, if any. Geographically remote 
records from outside the European region, such as those of Gurney 



(1927b) from the Suez Canal and Samoa, or those of Thomson 
(1882) (as Thorellia brunnea var. antarctica) from New Zealand, 
require verification. 

The body size of the Euryte longicauda from Mallorca falls in the 
range characteristic of the variety E. longicauda var. minor Scott, 
1905, which was elevated, rather inconsistently, to full specific 



98 



D. JAUME AND G.A. BOXSHALL 



status by Sars (1919-1921). This variety is supposed to inhabit 
deeper waters and never to occur in the littoral zone (Sars, 1913- 
1918; 1919-1921). 

According to Sars (191 3-191 8) £. longicauda carries only modi- 
fied flanged spines on the inner margin of the third exopodal 
segment of leg 2. In our material the proximalmost element on the 
inner margin of this segment is a plumose seta (Fig. 10B). However, 
this may be an observational error by Sars since material of E. 
longicauda from Raunefjorden in Norway (BMNH 1986.387) and 
from Scotland (BMNH 1951.8.10.587) in the collections of the 
Natural History Museum possesses a plumose seta in this position, 
as in the Mallorcan material. 



REVIEW OF EURYTE SPECIES 

The eight remaining species of Euryte are briefly reviewed here, in 
order to facilitate the identification of representatives of this prob- 
lematic genus. The review is essentially comparative and emphasises 
the most robust and reliable characters available in published de- 
scriptions. 

E. curticornis Sars, 1913 is characterised by short, 21 -segmented 
female antennules and the shortened third segment of the maxilliped; 
the curved distal claws on this appendage are also reduced in size 
and subsimilar in length. These features contrast with the maxilliped 
of E. longicauda, which is provided with an elongate third segment 
and with long, unequal distal claws. The distal spines on the third 
segment of endopod of leg 1 are clearly unequal in length in E. 
curticornis, whereas in E. longicauda they are about equal. 

E. longicauda can be distinguished from E. grata Herbst, 1989 
and E. verecunda Humes, 1992 by some features of the maxilla and 
maxilliped. In E. verecunda, the proximal spine on the distal coxal 
endite of maxilla is adorned on both sides with slender spinules. On 
the maxilliped, the armature element on the proximal syncoxal 
endite is a seta in E. longicauda, whereas in E. grata and E. 
verecunda this endite is represented by a stout spine. E. verecunda 
differs additionally in the setose condition of the armature elements 
on the inner margin of the second endopodal segment of leg 4; these 
elements are flanged spines in£. longicauda. The generic placement 
of E. verecunda needs verification since, according to Humes (1992), 
this species displays a 2-segmented condition of leg 5. This is a 
characteristic of the genus Ancheuryte Herbst, 1989, whereas in 
Euryte leg 5 is 3-segmented in both sexes. 

In E. pseudorobustaVervoort, 1964 two distal setae are present in 
the outer margin of the antennary coxobasis, whereas there is only 1 
seta in E. longicauda. The proximal spine on the distal coxal endite 
of maxilla has a different armature in the two species, with a row of 
setules along each side inf. pseudorobusta. Finally, the caudal rami 
of E. pseudorobusta are short, about as long as the anal segment, and 
differ significantly from the elongate caudal rami off. longicauda. 

Two other species, each described from a single female from the 
Addu Atoll (Maldives), viz. E. brevicauda Sewell, 1949 and E. 
sewelli Vervoort, 1964 (= 'Euryte sp.' of Sewell, 1949) also differ 
from E. longicauda in their very short caudal rami. The status of E. 
sewelli Vervoort, 1964 as a distinct species from E. brevicauda is 
equivocal (Vervoort, 1964; Sewell 1949). The main difference be- 
tween them is the apparently 18-segmented female antennule in the 
former species. Unfortunately, Sewell's (1949) original material is 
not preserved, thus precluding verification. However, if Sewell's 
illustrations are accurate, the 18-segmented antennule, combined 
with the absence of the inner seta on the proximal segment of exopod 
of leg 1 , can be used as diagnostic characters of this taxon. 



E. longicauda differs from E. bellatula Humes, 1 99 1 in the nature 
of the two armature elements on the inner margin of the second 
endopodal segment of leg 4; these are flanged spines in the former 
species, whereas in the latter they are setae. E. bellatula also has the 
proximal spine on the distal coxal endite of maxilla armed with a row 
of thin spinules on both sides. As commented above for£. verecunda, 
the generic placement of E. bellatula must be confirmed due to the 
apparently 2-segmented condition of the leg 5. The association with 
corals of the two taxa described by Humes is similar to the life-style 
of Ancheuryte, a closely related genus characterized by its 2-seg- 
mented leg 5. 

The status of E. similis Scott, 1912, originally described from the 
South Orkneys and never found since, is debatable. Scott pointed out 
its similarity to E. robusta, and that it appeared '. . . to differ in one 
or two minor points, such as in the armature of the first and fourth 
pairs of thoracic legs and in the proportional lengths of the abdomi- 
nal segments' (Scott, 1912). The differences in the armature of the 
swimming legs mentioned by Scott in the text do not correspond 
with his figures. Also, as Sewell ( 1 949) already pointed out, it seems 
certain that Scott had confused the legs so that his second leg is in 
reality the fourth, and his fourth leg is either the second or third. In 
fact, the original description is very superficial and does not permit 
any conclusion other than that the taxon belongs to Euryte. The only 
apparent diagnostic features displayed by this taxon could be the 
lack of an inner seta on both the first endopodal and first exopodal 
segments of leg 4 (Scott's leg 2). This is unreliable, however, since 
the number of armature elements on the swimming legs is a very 
conservative character at the generic level in the Cyclopidae. In our 
opinion, given the lack of type material, this taxon should be 
considered species inquirendum. 



Acknowledgements. We want to thank J. Pons-Moya and G. Pons their 
support during fieldwork. and Dr Rony Huys and Prof Carlos Eduardo 
Falavigna da Rocha for their detailed comments and improvements to the 
manuscript. Contribution to project DGICYT PB9 1-0055 and EC Training 
Research Contract ERBCHBICT941306. 



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Bull. not. Hist. Mm. bond. (Zool.) 62(2): 101-132 



Issued 28 November 1996 



Studies on the deep-sea Protobranchia 
(Rival via): the family Neilonellidae and the 
family Nuculanidae. 



J.A. ALLEN 

University Marine Biological Station, Millport, Isle of Cumbrae, Scotland, KA28 OEG 

H.L. SANDERS 

The Woods Hole Oceanographic Institution, Massachusetts, 02543, USA 

CONTENTS 

Synopsis 101 

Introduction 101 

Family Neilonellidae 102 

Neilonella salicensis (Seguenza 1877) 103 

Neilonella hampsoni (new species) 107 

Neilonella corpulenta (Dall 1881) 108 

Neilonella whoii (new species) 108 

Family Nuculanidae 1 1 1 

Subfamily Ledellinae 1 1 1 

Ledella acinula (Dall 1890) Ill 

Ledella aberrata (new species) 1 14 

Tmdariopsis agatheda (Dall 1889) 116 

Tmdariopsis aeolata (Dall 1890) 119 

Subfamily Nuculaninae 120 

Nuculana acuta (Conrad 1831 ) 120 

Nuculana commutata (Philippi 1 844) 123 

Nuculana vestita (Locard, 1898) 124 

Propeleda carpenteri (Dall 1881) 125 

Propeleda louiseae (Clarke 1961) 127 

Propeleda paucistriata (new species) 128 

Discussion 130 

References 131 



Synopsis. The morphology and distribution of four deep-water protobranch species of the family Neilonellidae and ten deep- 
water protobranch species of the family Nuculanidae are described. These include four new species. The evolution of the 
nuculanoid siphonate form is discussed. 



INTRODUCTION 



This is the ninth paper in a series on the deep-sea species of 
Protobranchia of the Atlantic. While a few undescribed species of 
families reported upon in earlier papers have been found in later 
samples and remain to be described, this paper is the last of our 
major descriptive accounts of the morphologies of this exceptionally 
important group of deep sea bivalves. 

In this paper we give an account of a number of nuculanacean 
species belonging to the families Neilonellidae and Nuculanidae 
present in our collections from the deep Atlantic. Related species 
already described are refered to only when essential for descriptive 
and comparative purposes. Unlike previous papers in this series 
most of the species described below are known, but only from their 
shell characters. This is in part due to the fact that many species come 
from upper-slope depths, and thus more likely to have been sampled 



in the past, and in part due to the fact that they are relatively large and 
thus less likely to have been lost due to the coarseness of nets used 
in earlier expeditions. Many earlier descriptions of the shell are far 
from adequate and new descriptions are given. 

We have followed the methods and approach of earlier papers in 
this series (Allen and Hannah, 1989; Allen and Sanders, 1973, 1982; 
Allen, Sanders and Hannah, 1995; Rhind and Allen, 1992; Sanders 
and Allen, 1973, 1977, 1985). From stereoscopic microscope ex- 
amination of shell features and whole mounts of the body stained in 
haemotoxylin, from dissected specimens and serial sections stained 
with trichome techniques, we give detailed descriptions of the shell 
and internal morphology of key species. Descriptions of related 
species are limited to points of difference and importance. 

Much of the material was taken by ourselves on numerous 
expeditions by research vessels of the Woods Hole Oceanographic 
Institution. These include the Atlantis', Atlantis II', 'Chain' and 
'Knorr'. Other samples were taken by JAA on the British research 



©The Natural History Museum. 1996 



102 



J.A. ALLEN AND H.L. SANDERS 



vessels 'Sarsia' and 'Discovery'. In addition to these, deep-sea 
bivalves from a series of French expeditions using the research 
vessels 'La Perle', 'Cryos' and 'Jean Charcot' were kindly donated 
by our French colleagues. At present this material, other than the 
types of new species and cited specimens, is housed at the Woods 
Hole Oceanographic Institution and the University Marine Biologi- 
cal Station Millport. On completion of our studies, it is our intention 
to return the French collections to the Museum National d'Histoire 
Naturelle, Paris and our own collections to the U.S. National Mu- 
seum and the Natural History Museum, London. 

Museums are abbreviated as follows:- ANSP - Academy of 
Natural Sciences, Philadelphia; BMNH - Natural History Mu- 
seum, London; MCZ - Museum of Comparative Zoology, Harvard; 
MNHN - Museum National d'Histoire Naturelle, Paris; SAM - 
South Africa Museum; USNM - United States National Museum, 
Washington, D.C.; ZMHU - Zoologisches Museum, Humbolt- 
Universitat, Berlin; ZMUC - Zoological Museum, University of 
Copenhagen. 

Sampling gear is abbreviated as follows:- AD - anchor dredge; CP 
- beam trawl; CV - Blake trawl; DS - epibenthic sledge Sanders 
(COB); ES - epibenthic sledge (WHOI); KG - USNEL boxcorer; 
OS - epibenthic sledge (SMBA); WS - epibenthic sledge (IOS). 



ABBREVIATIONS TO THE TEXT-FIGURES 



SR siphonal ridge 

SS style sac 

ST sensory tentacle 

SM stomach 

VG visceral ganglion 



Family Neilonellidae Schileyko 1989 

Shell usually robust, inflated, moderately large, not particularly 
elongate, more or less ovate, shell posteriorly extended to some 
degree and may be acutely rounded or carinate, usually marked 
concentric sculpture; umbo prominant, anterior to midline; hinge 
teeth taxodont, continuous or interrupted below the umbo; ligament 
external, sometimes with an internal component; siphons present; 
hind gut either a single loop or a series of loops and coils to the left 
and right of the body. 

The family comprises four genera, Neilonella, Pseudotindaria, 
Neilo and Pwtonucida (Allen and Hannah, 1986). By reason of their 
external ligament, the neilonellids in the past have been included in 
the family Malletiidae, however this family was shown to be an 
assemblage of unrelated taxa and is now much restricted (Sanders 
and Allen, 1985). The robust, ovate, concentically ridged shells of 
the neilonellids are in marked contrast to the thin, translucent, 
elongate shells, lacking any marked concentric sculpture, of the 
malletiids. 



AA 


anterior adductor muscle 


AN 


anus 


AR 


anterior pedal retractor muscle 


AS 


anterior sense organ 


BG 


'byssal' gland 


CG 


cerebral ganglion 


CM 


caecum 


CV 


cerebro-visceral commissure 


DD 


digestive diverticula 


DH 


dorsal hood 


DT 


digestive ducts 


FA 


feeding aperture 


FG 


fused gill tissue 


FR 


faecal rod 


FT 


foot 


GA 


gill axis 


GI 


gill 


GP 


gill plate 


GS 


gastric sheild 


HG 


hind gut 


HP 


hinge plate 


HT 


hinge teeth 


LI 


internal ligament or resilium 


LO 


opisthodetic external ligament 


MG 


combined mid gut 


MM 


mantle margin 


OE 


oesophagus 


PA 


posterior adductor muscle 


PE 


sensory papilla 


PG 


pedal ganglion 


PL 


palp 


PP 


palp proboscis 


PR 


posterior pedal retractor muscle 


PS 


posterior sorting area 


SE 


siphonal embayment 


SP 


combined siphon 



Genus NEILONELLA Dall 1881 

TYPE SPECIES. Leda (Neilonella) corpulenta Dall 1881; OD. 

Shell moderately robust, relatively wide, strong concentric sculp- 
ture, anterior margin rounded, posterior margin slightly attenuate 
and may be acutely rounded, not carinate; umbo prominant, anterior 
to the midline; hinge plate moderately strong, many chevron teeth, 
small gap between anterior and posterior series; external ligament 
opisthodetic, inserting in an external gutter in each valve, with 
minute internal resilifer, siphonal embayment shallow; hind gut 
single loop to the right of the body. 



Genus PSEUDOTINDARIA Sanders and Allen 1977 

TYPE SPECIES. Pseudotindaria erebus (Clarke 1959); OD. 

Shell robust, relatively wide, anteriorly and posteriorly rounded, 
not carinate, strong concentric sculpture; hinge plate strong with 
numerous chevron teeth continuous below umbo; ligament external, 
anterior part short, posterior part elongate; hind gut with complex 
series of loops and coils to the right and left of the body. 

Genus NEILO Adams 1854 



Type species. Neilo cumingii Adams 1854; by monotypy. 

Shell robust, relatively wide, moderately elongate and posteriorly 
extended, postero-dorsal margin straight or slightly concave, cari- 
nate, posteriorly truncate or slightly rostrate, postero- ventral margin 
may be somewhat sinuous, two rounded radial ridges from umbo to 
posterior margin, concentric sculpture; hinge plate well-developed; 
numerous chevron teeth, anterior and posterior series separate and 
may differ in size (anterior larger); ligament external. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 
Genus PROTONUCULA Cotton 1930 

Type species. Protonucula verconis Cotton 1930; OD. 

Shell relatively fragile, oval, compressed, glossy, concentric sculp- 
ture; umbo prominent; hinge plate narrow, chevron teeth squat, 
broad, continuous; ligament relatively small, external, amphidetic. 

Neilonella salicensis (Seguenza 1877) 

Type specimen. Whereabouts unknown. 

Type locality. Upper Pliocene fossil, southern Italy. 

Cited specimens. BMNH 1995051. 

Leda pusio var./af/or Jeffreys 1876a, p. 190, nom. nud.; 1876b, 

p.430. 
Leda pusio Jeffreys 1879, p. 578, in part. 
Leda pusio var. salicensis Seguenza 1877, p. 1 178, pi. 4, fig. 20. 
Yoldia sericea var.striolata Verrill 1884, p.226; 1885, p.576. 
Leda salicensis Locard 1898, p.348, pi. 14. figs 22-25. 
Neilonella subovata Verrill and Bush 1 897, p.57, figs 7, 8. 22; type 

locality: Georges Bank to Cape Hatteras, 1 25-1 73 1 fms; syntype: 

USNM 34826 (examined by HLS). 
Tindaria (Neilonella) guineensis Thiele 1 93 1 , in Theile and Jaeckel 

1931, p.50, pi. 3, fig.70; type locality; Gulf of Guinea, 710- 

2492m; type: ZMHU (not seen). 
Neilonella guineensis Knudsen 1970, p.59, text-fig. 39, pi. 5, fig. 3, 

pi. 6, figs 8-10. 
Neilonella schepmani Prashad 1932, p. 27, pl.l, figs 50, 51; type 

locality; Banda Sea, Siboga sta.221, 2798m; ZMHU (not seen). 
Pseudoneilonella salicensis Laghi 1986, p. 191, text fig.l, pi. 5, figs 

1-7. 
Pseudoneilonella latior Laghi 1986, p. 192, text fig. 1 , pi. 7, figs 7a-c. 
Pseudoneilonella salicensis atlantica Laghi 1986 p. 192, text fig.l, 

pi. 6, figs 1-7. 
Pseudoneilonella montanaroe Laghi 1986, p. 193, text fig.l, pl.9, 

figs 4-8; type locality: off Portugal, Porcupine Expedition, 400m, 

USNM No. 199739 (not seen). 
Neilonella striolata (Brugnone) Waren 1989, p. 252, figs 3a, 16a-d. 

Waren (1989) gives a lengthy and detailed discussion of much of 
the nomenclatoral history of this species although, the history is 
even more complex than he reports. Waren (1989) synonymized N. 
salicensis with an earlier described species N. striolata (Brugnone, 
1876). The type of AA striolata is lost but from the original descrip- 
tion and figure by Brugnone (1876) and those of Seguenza (1877) 
(who synonymized it with Yoldia abyssicola Torrell) and later au- 
thorities (e.g. Jeffreys, 1879; Locard, 1898) it is clear that the 
posterior part of the shell of N. striolata is significantly different in 
form to that of N. salicensis. Whereas the posterior shell margin of 
N. salicensis is acutely rounded that of N. striolata is sub-rostrate 
with the posterior limit of the shell more ventral in position and the 
distal postero-dorsal margin more steeply angled (Figs 1,2 & 4). 

The brief description of N. guineensis by Theile in Theile and 
Jaeckel( 1 93 1 ) was enlarged upon by Knudsen (1970) who com- 
pared specimens from the Galathea Expedition with those of the 
Valdivia and Siboga Expeditions. From our examination of the 
Galathea material from 02"17'S 08T0'E (2770m) and our own speci- 
mens from 02"32'S 08'18'E (2514m) we conclude that N. guineensis 
and N. salicensis are the same species. Knudsen (1970) also 
synonymized N. schepmani with N. guineensis. Although N. 
schepmani is from the Banda Sea (Prashad, 1932) which might cast 
doubt on this synonymy, it must be remembered that N. salicensis is 
a very widespread species and we believe that it is comparable to 



103 




Fig. 1. 'Yoldia abyssicola' Torell. Right valve, redrawn from Seguenza 
(1877), synonymized with Leda producta Monterosato and Yoldia 
striolata Brugnone. Scale = 1mm. 

Ledella sublevis, which also has a widespread distribution in the 
Atlantic and extends into the southwest Pacific (Allen and Hannah, 
1989). 

Further compexity in the synonymy ofN. salicensis concerns so- 
called varieties of "Leda pusio Philippi' a species that has recently 
been investigated by Laghi (1986). 




Fig. 2. Neilonella salicensis. a, view of shell, from Atlantis II station 73 
in the North America Basin, drawn from the left side; b, outline of shell 
from Biogas VI station DS 86 in the West European Basin to show 
difference in shape. Scale = 1mm. 

L. pusio is rostrate and not a neillonellid, but a ledellid, possibly 
synonymous withL.acuminata (Jeffreys 1 870)(Laghi, 1986). Jeffreys 
( 1 879) reporting on Leda pusio taken by the 'Lightning' and 'Porcu- 
pine' Expeditions, refers to what he thought was a variety of this 
species which he had previously named L. pusio var. latior 
(Jeffreys, 1876) and which he changed to L. pusio var.salicensis of 
Seguenza (1877). Laghi (1986) proposed a new genus 
Pseudoneilonella to accommodate this latter and raised to species 
level a number of the records of Jeffreys ( 1 876, 1 879). Since then, 
Waren (1989) has synonymized these with Neilonella salicensis and 
this we confirm. 

Jeffreys (1879) also listed a further variety which he called 
semistriata and which is now regarded as a separate species (Waren, 
1989). In his detailed analysis Waren (1989, figs 17E & F) also 



104 



J.A. ALLEN AND H.L. SANDERS 



figures what he describes as 'Neilonella latior (Jeffreys)?, young 
syntype of Leda sericea Jeffreys, Valorous Expedition, Stn 12, 
USNM 199595' These valves are equilateral in shape and differ in 
outline from N. salicensis. We have specimens of this species in our 
collections and these we intend to describe in a later paper. 

Locard ( 1 898), not mentioned by Waren ( 1 989), came to the same 
conclusion as ourselves and recognized three entities, Leda pusio, 
L. striolata and L. salicensis, accurately separating them on shell 
characters, the most significant of these being the more elongate, 
triangular form and greater post-umbonal length of L. salicensis as 
compared with L. pusio, both of which he figures. 

There is a relatively wide degree of variation in the height, length 
and post-umbonal length of the shell in N. salicensis and this 
variation is similar in populations from different basins (Figs 4 & 5). 
This may, in part, explain why this species has been described anew 
so many times. 

Material: 

Cruise Sta Depth No Lat Long Date Gear 

(m) 



NORTH AMERICA BASIN 



AtlantisII 


62 


2496 


13 


39"26.0'N 


70°33.0'W 


21.8.64 ES 


12 


64 


2886 


2 


38"46.0'N 


70"06.0'W 


21.8.64 ES 




73 


1330- 

1470 


495 


39"46.5'N 


70°43.3'W 


25.8.64 ES 


Chain 50 


87 


1102 


6 


39'48.7'N 


7040.8'W 


6.7.65 ES 


Chain 58 


103 


2022 


217 


3943.6'N 


70'37.4'W 


4.5.66 ES 


AtlantisII 30 131 


2178 


119 


3938.5'N 


70°36.5'W 


18.12.66 ES 


Chain 88 


210 


2024- 
2064 


48 


39'43.0'N 


70°55.5'W 


23.2.69 ES 


GUYANA BASIN 












Knorr 25 


293 


1456- 
1518 


2 


08°58.0'N 


54Tj4.3'W 


27.2.72 ES 




295 


1000- 
1022 


1575 


08"04.2'N 


54"21.3'W 


28.2.72 ES 




297 


508- 

523 


194 


07°45.3'N 


54°24.0'W 


28.2.72 ES 




299 


1942- 
2076 


1 


0755. l'N 


55"42.0'W 


29.2.72 ES 




307 


3835- 
3862 


2 


12°34.4'N 


58"59.3'W 


3.3.72 ES 


WEST EUROPEAN BASIN 








Sarsia 


S33 


1537- 
1830 


6 


43°41.0'N 


03°36.0'W 


13.7.67 ES 




S37 


1739 


2 


43°39.2'N 


03°30.2'W 


15.7.67 ES 




S40 


860 


4 


43"35.6'N 


03"24.8'W 


15.7.67 ES 




S44 


1739 


60 


43"40.8'N 


03"35.2'W 


16.7.67 ES 




S50 


2379 


16 


43'46.7'N 


03"38.0'W 


18.7.67 ES 




S65 


1922 


2 


46°15.0'N 


04°50.0'W 


25.7.67 ES 


Chain 106 


313 


1491- 
1500 


432 


51°32.2'N 


12'35.9'W 


17.8.72 ES 




316 


2173- 
2209 


583 


50"58.7'N 


13'01.6'W 


18.8.72 ES 


La Perle 


DS11 


2205 


1 


47"35.5'N 


08°33.7'W 


8.8.72 DS 


Biogas I 














J.Charcot 


DS25 


2096 


9 


44°08.2'N 


04°15.0'W 


1.11.72 DS 


Polygas 














J.Charcot 


DS32 2: 


2 


4732.2'N 


08°05.3'W 


19.4.73 DS 


Biogas II 














BiogasIII 


DS38 


2138 


2 


47'32.2'N 


08°35.8'W 


25.8.73 DS 




DS49 


1845 


70 


44°05.9'N 


04°15.6'W 


31.8.73 DS 


Biogas IV 


DS52 


2006 


16 


44'06.3'N 


04"22.4'W 


18.2.74 DS 




DS63 


2126 


5 


4732.8'N 


08"35.0'W 


26.2.74 DS 


Cryos 


CP07 


2170 


5 


44°09.8'N 


04'16.4'W 


21.6.74 CP 


Biogas V 














J.Charcot 


DS77 


4240 


1 


47°31.8'N 


09'34.6'W 


24.10.74 DS 


Biogas VI 


DS80 


4720 


4 


46'29.5'N 


10°29.5'W 


26.1 0.74 DS 




DS82 


4462 


1 


44°25.4'N 


04"50.8'W 


29.10.74 DS 




DS86 


1950 


198 


44'04.8'N 


04"18.7'W 


31.10.74 DS 





DS87 


1913 


173 


44°05.2'N 


04°19.4'W 


1.11.74 


DS 




DS88 


1894 


31 


44°05.2'N 


04'15.7'W 


1.11.74 


DS 


Incal 


DS01 


2091 


98 


5759.2'N 


10°41.3'W 


15.7.76 


DS 




DS02 


2081 


91 


5758.2'N 


10"48.5'W 


16.7.76 


DS 




CP01 


2068 


19 


5757.7'N 


10°55.0'W 


16.7.76 


CP 




DS05 


2503 


3 


56'28.1'N 


11*11.7'W 


18.7.76 


DS 




OS01 


2634 


4 


50'14.4'N 


13"10.9'W 


30.7.76 


OS 




WS02 2498 


1 


50°19.3'N 


12°55.8'W 


30.7.76 


WS 




CP10 


4823 


2 


48°25.5'N 


15"10.7'W 


31.7.76 


CP 




WS03 4829 


1 


48"19.2'N 


1513.3'W 


1.8.76 


WS 


Challenger 


232 


2195 


3 


5717.0'N 


10°16.0'W 


19.5.83 


ES 


7/83 
















CANARY BASIN 














Discovery 


6701 


1934 


8 


2745.2'N 


14°13.0'W 


16.3.68 


ES 


SIERRE LEONE BASIN 












AtlantisII 


142 


1624- 


20 


10"30.0'N 


17'51.5'W 


5.2.67 


ES 


31 




1796 














147 


2934 


16 


10°38.0'N 


17'52.0'W 


23.2.67 


ES 


ANGOLA BASIN 














AtlantisII 


191 


1546- 


52 


23°05.0'S 


1731.5'E 


17.5.68 


ES 


42 




1559 














200 


2644- 
2754 


7 


09"43.5'S 


10"57.0'E 


22.5.68 


ES 




201 


1964- 
2031 


41 


09"25.0'S 


11'35.0'E 


23.5.68 


ES 




202 


1427- 
1643 


21 


08'56.0'S 


1715.0'E 


15.5.68 


ES 


Walda 


DS20 


2514 


4 


02"32.0'S 


08"18.1'E 


-.-.71 


DS 




DS25 


2470 


31 


0ri9.8'S 


0749.2'E 


-.-.71 


DS 



In addition to the material listed above we have examined the 
following specimens from the Porcupine and Valorous Expeditions 
housed in the BMNH:- 85. 1 1 .5.397^02; 6.9.27,28; 85. 1 1 .5.494-5; 
89.1 1.1 1.10-13; 89.9.5.26-9; 77.1 1.28.24. We have also examined 
specimens from the Galathea Expedition housed in the ZMUC, and 
specimens mostly from the U.S. Fish Commission expeditions housed 
in the USNM and MCZ. 

Distribution. Common throughout the Atlantic north of the 
Argentine and Cape Basins and south of the Norwegian and Arctic 
Basins. It is also found in the southwestern Pacific. It mainly occurs 
at slope depths, occasionally at abyssal depths (West European 
Basin), from 508m to 4829m. 




Fig. 3. Neilonella salicensis. a, internal inclined ventral view of the 
hinge plate and teeth of a left valve; b, dorsal view of shell to show the 
extent of the external ligament and detail of sculpture. Both specimens 
taken from Walda station DS 25 in the Angola Basin. Scale = 1mm. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



105 




Fig. 4. Neilonella salicensis. A series of shells, from Atlantis II station 73 
from the North Atlantic Basin, in lateral view from right side to show 
change in shape with increasing size. Scale = 1mm. 

Shell description (Figs 2-A) 

Shell robust, moderately elongate, moderately wide, ornamented 



with concentric ridges sometimes in groups of three to four well- 
defined ridges alternating with two to to three slighty less 
well-defined ridges slightly wider spaced than the former, light to 
dark straw-coloured periostracum; umbos moderately prominant, 
somewhat anterior to mid-line (postumbonal length c.54% of total 
length), inward and slightly anterior facing; dorsal shell margins 
deep set close to umbo but no lunule or true escutcheon, antero- 
dorsal margin slopes down to join anterior and ventral margins in 
smooth curve, postero-dorsal margin almost straight, may be 
slightly angled at posterior limit of hinge short of acutely rounded 
posterior margin; posterior limit of shell somewhat dorsal to mid 
horizontal line, anterior limit of shell at mid line; posterior margin 
of smaller specimens less acutely rounded; hinge plates elongate, 
moderately broad, increasing slightly in width distally, anterior 
and posterior hinge teeth series meet below umbo with no edentu- 
lous space between, teeth numerous, up to 19 in posterior series 
and 14 in anterior series depending on size of specimen, obtuse 
chevron-shape, proximal hinge teeth very small, those more distal 
elongate and so closely articulated that it maybe impossible to 
separate valves without damage to them; internal ligament 
(resilium) microscopic, present ventral to umbo at outer margin of 
hinge plate, external ligament opisthodetic, relatively short, short 
anterior part present. Maximum total length of present specimens 
8.0mm. 

As in most deep-sea protobranchs, there is a change in the shape 
of the shell outline as growth occurs (Figs 4 & 5), although this is not 
as marked as in many protobranch species (e.g. Ledella spp. Allen 
and Hannah, 1989). The ratio of postero-umbonal length to total 
length increases and at the same time the height to total length 
decreases so that larger shells (>3mm total length) are more 
posteriorly attenuate than smaller specimens. There is a fair degree 
of variation in the ratios which is not related to the different 
populations sampled. Unusually in some samples the length fre- 
quency histograms are skewed to the right (Fig. 6) and with clearly 



ii o 70-J 
TL S 



60 - 



PL , 
TL " 



50 




Length (mm) 

Fig. 5. Neilonella salicensis. The ratios height to total length (H/TL)(large symbols) and post-umbonal length to total length (PL/TL)(small symbols) 
plotted against total length to show changes in shell proportions with increasing length. Open circles are specimens from Biogas III station DS 49 West 
European Basin, closed circles from Sarsia Station S44 West European Basin and closed squares, from Chain 58 station 103 North America Basin. 



106 



J.A. ALLEN AND H.L. SANDERS 




LENGTH (mm) 

Fig. 6. Neilonella salicensis. Length/frequency histogram of a sample taken from Atlantis II station 73 from the North America Basin. 



marked size groups. It is unclear whether these latter relate to annual 
settlements or to fortuitous periodic settlements. 

Internal morphology (Figs 7 & 8) 

This was one of the few deep-sea species that were kept alive for a 
few weeks following the processing of the sample. The external drab 
olive/yellow of the shell is in marked contrast to the spectacularly 
vivid array of colours of the body organs. The stomach is a bright 
emerald green, the digestive gland a bright olive green and the gills 
are bright red. The gill plates are particularly vivid below the frontal 
cilia and dorso-laterally, in addition the mantle overlying the gills is 
also red. The margins of the palps are a diffuse pink as too is a band 
underlying the crest of the inner folds. The red and pink pigments are 
probably a cytochrome. The blood is not red in colour and probably 
contains a haemocyanin as do other protobranchs (Taylor, Daven- 
port and Allen, 1995). Elsewhere the palps are pale yellow and the 
yellow/pink effect is in marked contrast to the palp proboscides 
which are pure white. 

Gl CV DP 




Fig. 7. Neilonella salicensis. Semidiagrammatic view of the internal 
morphology as seen from the right side. See abbreviations to text-figures 
on p. 102. Scale = 1mm. 



The mantle has three marginal folds of which the inner is fused at 
the posterior end to form the siphon. Fusion is minimal, restricted to 
the dorsal margin of the siphon (Fig. 8). Ventrally the siphonal 
margins are unfused as too are the longitudinal ridges that mark the 
division between the dorsal exhalent and ventral inhalent channels. 
The gill axes attach to the inner ends of the longitudinal ridges. 
There is a shallow siphonal embayment and attached to its inner 
limit on the left or the right side is a siphonal tentacle which in living 
specimens extends beyond the shell margin at distance equal to a 
third of the shell length. Ventral to the anterior adductor muscle there 
is an elongate anterior sense organ, which consists of a flap of tissue 
derived from the middle sensory fold. A major ciliated rejection tract 
is present on the inner face of the inner mantle fold. The adductor 
muscles are relatively small, unequal in size, with the posterior the 
smaller. The posterior muscle is round in cross-section while the 
anterior is bean-shaped. 

The gills, with up to 26 plates, are slung from a horizontal axis 
extending from the base of the siphon to a point about half way 
across the body. Individual gill plates are exceptionally elongate for 
a deep-sea protobranch. The foot is broad and the divided sole has 
papillate margins, the anterior papillae are the more pronounced. 
There is a small median papilla present posterior to the aperture to 
the 'byssaF gland. The latter is extremely large. The pedal muscles 
consist of a pair of broad posterior retractors, one on each side of the 
hindgut anterior to the posterior adductor, and a series of four pairs 
of anterior retractors posterior to the anterior adductor. The two 
inner muscles of the four lie more close together than to the two 
outer. 

The palps are relatively large with up to 27 inner ridges depending 
on the size of the specimen. Each bears a long narrow palp probos- 
cis. A ciliated rejection tract traverses the lateral face of the foot at 
the junction of of the muscular and visceral parts and the posterior 
ventral point of the palp is positioned at the posterior limit of this 
tract in the living animal. 

The mouth is set far posterior to the anterior adductor. The 
oesophagus curves first antero-dorsally to meet the inner face of the 
anterior adductor then postero-dorsally to open on the anterior face 
of the stomach. The stomach is large and slung diagonally within the 
visceral mass. The pedal muscles form a ventral 'basket' in which 
the stomach rests. The stomach, which is similar to that of other 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



107 





Fig. 8. Neilonella salicensis. Detail of internal morphology; a, lateral 
view from the right side of the stomach and style sac; b, internal view of 
a dissected gastric shield; c, diagrammatic enlargement of the siphonal 
region to show the relationship of the parts. See abbreviations to the 
text-figures p. 102. Scales = 0.5mm (A); 0.1mm (B). 

protobranchs (Purchon, 1956), is one third lined with agastric shield 
dorsally and to the left (Fig. 8). The unlined right wall bears very few 
sorting ridges. The style sac is flask-shaped rather than the more 
usual conical outline of other protobranchs. The combined mid gut 
is guarded by minor and major typhlosoles. The hind gut penetrates 
the foot to a point ventral to the pedal ganglia before taking a course 
posterior to the stomach and thence to form a loop to the right side 
of the body. This single loop extends anteriorly as far as the anterior 
adductor muscle and where it then takes a dorsal course along the 
margin of the body to the anus. The hind gut has a particularly wide 
diameter compared with most other deep-sea protobranchs and has 
a well-marked typhlosole present along its entire length. The diges- 
tive gland which lies anterior and dorso-lateral to the stomach has 
three sections each connecting with the stomach via a duct. The 
apertures of the ducts are ventral, and left latero-ventral to the 
oesophageal aperture. 

The nervous system is of a typical protobranch design with 
elongate pairs of cerebral and visceral ganglia and large ovoid pedal 
ganglia. These are connected by very stout cerebro-visceral and 
cerebro-pedal commissures. A statocyst lies dorsal each pedal gan- 



glion. From visceral ganglia three main nerves pass to the gill, hind 
gut and mantle/siphon, and from the cerebral ganglia nerves pass to 
the palps and mantle. 

The sexes are separate, specimens greater than 4mm total length 
have gonads. The gonad first develops at the periphery of the visceral 
mass and as it matures it spreads across the lateral surface of the 
viscera. On the right side of the body, apart from a small portion 
postero-dorsal to the loop of the hind gut the gonad does not extend 
beyond the confines of the of the loop. While different samples show 
different degrees of maturity, the specimens of each sample appear 
to be maturing synchronously. The ratio of males to females is 
approximately 50:50. 

Neilonella hampsoni new species 

Type SPECIMEN. Holotype BMNH 1995049; paratype BMNH 
1995050. 

Type LOCALITY. Atlantis II cruise 31, station 155, Sierra Leone 
Basin, 00°03.0'S 27°48.0'W, 3730-3783m. 

Material: 

Cruise Sta Depth No Lat Long Date Gear 

(m) 

SIERRA LEONE BASIN 

Atlantis 11 155 3730-3 00°03.0'S 27°48.0'W 14.2.67 ES 

31 3783 

J.Charcot DS03 5150 1 10°59.0'N 45°15.0'W 16.1 1.77 DS 

Vema 



Distribution. An abyssal species, to date only found in the Sierra 
Leone Basin at depths from 3730m to 5150m. 

Shell description (Figs 9 & 10) 

Shell robust, oval, wide, ornamented with concentric ridges, straw- 
coloured periostracum; umbo prominant, anterior to midline 
(postumbonal length c.60% of total length in large specimens), 
inward facing; lunule and escutcheon, both well-defined; postero- 
dorsal margin in lateral view almost straight, slopes relatively 
steeply from umbo to proximal limit of hinge, rounded angle to 
posterior margin, anterior and ventral margins form a smooth curve, 
distally antero-dorsal margin slightly concave then slopes steeply in 
smooth curve to anterior margin; anterior limit of shell coincides 




Fig. 9. Neilonella hampsoni. Lateral view of the right valve of the 
holotype, from Atlantis II station 155 from the Sierra Leone Basin. 
Scale = 1mm. 



108 



J.A. ALLEN AND H.L. SANDERS 




Fig. 10. Neilonella hampsoni. a, lateral view of the hinge plate of the 
right valve of a paratype; b, dorsal view of the shell of the holotype. 
Both specimens taken from Atlantis II station 155 from the Sierra Leone 
Basin. Scales = 1mm. 

with horizontal midline, posterior limit of shell immediately ventral 
to mid line; hinge plate stout extending along most of the dorsal shell 
margin, anterior and posterior tooth series meet below umbo, with 
faint suggestion of an edentulous space between, 1 3 posterior and 1 
anterior teeth in specimen 3.9mm total length, teeth increase in size 
distally, teeth chevron-shaped, obtuse, so much so that teeth appear 
to be a straight line transverse to hinge plate; ligament opisthodetic, 
external, short, anterior outer layer extremely short, hidden beneath 
umbo, no resilium. 

Maximum total length of present specimens is 7.5mm. 

Apart from N. salicensis, from which it differs in having a shorter, 
wider shape, and well-marked lunule and escutcheon, the only other 
protobranch species with which it has some similarity is 'Leda' 
sericeavar ovata Jeffreys 1 876 (Jeffreys, 1 879). One of us (JAA) has 
examined specimens of this latter species in the Natural History 
Museum, London, (BMNH 85 1 1 5483-84) and find that L. sericea 
is more ovate, with the dorsal margins much less sloping, a more 
anteriorly positioned umbo and a much more narrow hinge plate. 

Internal morphology 

The morphology is similar to Neilonella salicensis. Such differences 
that do exist include the adductor muscles, both of which are small, 
similar in size, but with the posterior oval and the anterior round in 
cross section. The foot is somewhat smaller but with a moderately 
large byssal gland with a small, hooked, median papilla posterior to 
its aperture. Except for the posterior quarter of their length, the 
margins of the divided foot are finely papillate. There are approxi- 
mately 20 gill plates and 25 ridges on the palps of a specimen 3.9mm 
total length. The hind gut makes a simple loop to the right side of the 
body, it has a wide lumen (0.21mm in diameter) with a single well- 
defined typhlosole running its entire length. The stomach is large 
and the mouth lies some distance posterior to the inner wall of the 
anterior adductor. The kidney extends in a narrow band from the 
posterior margin of the posterior adductor, anteriorly narrowing over 
the viscera, and terminating at the posterior edge of the digestive 
diverticula. 



It is named after our good friend and colleague George Hampson 
who accompanied us on so many of our cruises and without whom 
sampling at abyssal depths would not have been the resounding 
success that it proved to be. 

Neilonella corpulenta (Dall 1881) 

TYPE LOCALITY. Blake station off Havana, 823m (station number 
not recorded but, only station 51, 23Tl.0'N 82°21.0"W, is listed as 
having a depth of 450fm (823m) (Smith, 1888)). 

Type specimen. Holotype, U.S. National Museum 63169. Cited 
specimen: BMNH 1995048. 

Leda (Neilonella) corpulentaDatt, 1881, 125; 1886, 254, pi. 7, figs. 

la, lb. 
Neilonella (N.) corpulenta Laghi 1986, pi. 9, figs 1-3. 



Material: 










Cruise Sta 


Depth No 
(m) 


Lat 


Long 


Date Gear 


BRAZIL BASIN 

Atlantis II 162 
31 


1493 1 


08'02.2'S 


34°03.0'W 


19.2.67 ES 



The type specimen has been examined by us. 

Distribution. An upper slope species previously recorded only 
from the Gulf of Mexico but here found in the Brazil Basin. It occurs 
at depths from 347m to 1493m. 

Shell description (Fig. 11) 

Dall (1881) gave an adequate description which was later (Dall, 
1 886) augmented by good internal and external lateral views of the 
shell. The specimen collected from the Brazil basin differs little 
from the type (Dall, 1881, 1886)(Fig. 11):- 

Shell elongate, solid, oval, ornamentated with concentric ridges; 
umbo not particularly large or elevated, somewhat anterior to the 
mid line; postero-dorsal margin almost straight, slightly upturned 
posterior to the distal limit of the hinge, then sharply and smoothly 
curved to posterior margin, ventral margin shallow smooth curve, 
not posteriorly sinuous, anterior margin smoothly curved, antero- 
dorsal margin relatively steeply angled, distally slightly raised; 
hinge plate elongate, relatively wide, hinge teeth chevron-shaped, 9 
in anterior and 12 in posterior series; external ligament slightly 
opisthodetic, short; resilium small ventral to umbo. 

Dall ( 1881 ) states that there are an equal number of teeth in the 
anterior and posterior series (15), however the type specimen which 
is larger than the present specimen, has 17 anterior and 20 posterior 
teeth. 

The total length of present specimen is 3.1mm. 

Internal morphology 

Both adductor muscles are oval in cross section, the anterior is 
somewhat larger than the posterior but neither is particulaly large. 
The foot is relatively short with large marginal papillae. The palps 
are relatively short with 7 broad internal ridges. The gill is also short. 
The siphonal embayment is shallow and the siphons similar to those 
described for N. salicensis. The hind gut forms a single loop to the 
right side of the body and has a typhlosole along its entire length. 

Neilonella whoii new species 



Type specimen. 
1995053. 



Holotype BMNH 1995052; paratypes BMNH 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



109 




Fig. 11. Neilonella corpulenta. a, an external lateral view of the right valve of specimen No 63169 from the USNM and an internal lateral view of the 
central region of the hinge of the same valve; b. lateral view of the intact shell, from Atlantis II station 162 from the Brazil Basin, and an internal view of 
the left valve of the same specimen to show detail of the hinge plate. Scales = 1 mm. 



Cited specimen. BMNH 1995054. 








Biogas IV 


DS53 4425 


30 


4430.4'N 


04"56.3'W 


19.2.74 


DS 


Type locality. Chain cruise 50, station 78, North America Ba- 
sin, 38°00.8'N 69'18.7'W, 3828m. 




DS54 4659 
DS56 4050 
DS60 3742 


13 

2 

2 


46'31.1'N 
4732.7'N 
47*26.8'N 


10'29.2'W 
09"28.2'W 
09Tj7.2'W 


21.2.74 
23.2.74 
24.2.74 


DS 
DS 
DS 


Material: 














Cryos 
Biogas V 


DS66 3480 


4 


4728.2'N 


09"00.0'W 


17.6.74 


DS 


Cruise 


Sta Depth 

(m) 


No 


Lat 


Long 


Date 


Gear 


J.Charcot 
Biogas VI 


DS75 3250 
DS76 4228 
CPU 4237 
DS78 4706 


4 
8 
1 
29 


47*28. l'N 
4734.8'N 
47°32.0'N 
46"31.2'N 


09"07.8'W 
09°33.3'W 
09'35.9'W 
10°23.8'W 


22.10.74 
23.10.74 
23.10.74 
25.10.74 


DS 
DS 

cp 


WEST EUROPEAN BASIN 










DS 


Chain 106 


323 3356- 
3338 


7 


50"08.3'N 


13'53.7'W 


21.8.72 


ES 




DS79 4715 
CP18 4721 


10 

1 


46"30.4'N 
46"30.5'N 


10°27.1'W 
10"26.0'W 


26.10.74 
26.10.74 


DS 
CP 




326 3859 


5 


50"04.9'N 


14'23.8'W 


22.8.72 


ES 




DS81 4715 


1 


46'28.3'N 


10°24.6'W 


27.10.74 


DS 




328 4426- 


6 


50"04.7'N 


15"44.8'W 


23.8.72 


ES 




CP19 4434 


3 


44°24.9'N 


0451. 3'W 


28.10.74 


CP 




4435 














DS82 4462 


16 


44"25.4'N 


04°52.8"W 


29.10.74 


DS 




330 4632 


16 


50°43.5'N 


IT51.TW 24.8.72 


ES 


J.Charcot 


OS01 2634 


3 


50"15.2'N 


13°11.0'W 


30.7.76 


OS 


J.Charcot 


DS20 4226 


2 


47'33.0'N 


09°36.7'W 


24.10.72 


DS 


Incal 


DS11 4823 


1 


48°18.6'N 


15'12.0'W 


1.8.76 


DS 


Poly gas 


DS22 4144 


2 


47°34.1'N 


09°38.4'W 


25.10.72 


DS 




CPU 4823 


1 


48'21.2'N 


15'13.7'W 


1.8.76 


CP 




DS23 4734 


18 


46"32.8'N 


10°21.0'W 


26.10.72 


DS 




OS02 4829 


6 


48°19.1'N 


15'15.5'W 


2.8.76 


OS 




DS28 4413 


1 


44"23.8'N 


04°47.5'W 


2.11.72 


DS 




OS05 4296 


3 


47'32.9'N 


09'34.7'W 


7.8.76 


OS 


J.Charcot 


CV23 2034 


1 


47°32.7'N 


08"34.2'W 


25.8.73 


CV 




OS06 4316 


3 


47'27.9'N 


09°36.0'W 


9.8.76 


OS 


Biogas III 


DS41 3548 


4 


47'28.3'N 


09"04.2'W 


26.8.73 


DS 




DS16 4268 


1 


47°30.3'N 


09°33.4'W 


9.8.76 


DS 




CV27 4023 


1 


4734.2'N 


09°32.4'W 


28.8.73 


CV 




WS094277 


2 


47"27.9'N 


09"34.0'W 


10.8.76 


WS 




CV30 4518 
DS47 4230 


53 

2 


46°32.8'N 
44"26.8'N 


10°20.0'W 
0450.7'W 


28.8.73 
31.8.73 


CV 
DS 


CANARY BASIN 












J.Charcot 


CV34 4406 


1 


44"27.2'N 


04°49.rW 


19.2.74 


CV 


Discovery 


6711 2988 


1 


2714.9'N 


15"36.3'W 


19.3.68 


ES 



110 



J.A. ALLEN AND H.L. SANDERS 



SIERRA LEONE BASIN 












Atlantis II 


146 


2842- 


2 


10'39.5'N 


17°44.5"W 


6.2.67 


ES 


31 




2891 














147 


2934 


4 


10"38.0'N 


1752.0'W 


6.2.67 


ES 




148 


3814- 4 


10°37.0'N 


18T4.CW 


7.2.67 


ES 






3828 














149 


3861 


1 


10"30.0'N 


18'18.0'W 


7.2.67 


ES 


GUINEA BASIN 














J. Charcot 


DS2S 


1261 


2 


04"21.2'N 


04°35.2'E 


7.8.71 


DS 


waifla 
ANGOLA BASIN 














Atlantis II 


195 


3797 


46 


14"49.0'S 


09'56.0'E 


19.5.68 


ES 


42 


197 


4595 


25 


10"29.0'S 


09TJ4.CE 


21.5.68 


ES 




198 


4559- 
4566 


20 


1C24.CS 


09"09.0'E 


21.5.68 


ES 




199 


3764- 
3779 


3 


09°49.0'S 


10"33.0'E 


22.5.68 


ES 




200 


2644- 
2754 


8 


09"43.5'S 


10°57.0'E 


22.5.68 


ES 


NORTH AMERICA BASIN 










Atlantis II 


2 


3752 


2 


38°05.0'N 


69"36.0'W 


22.5.61 


AD 


264 
















Atlantis II 


64 


2886 


2 


38"46.0'N 


70'06.0'W 


21.8.64 


ES 


12 


72 


2864 


9 


38T6.CN 


71"47.0'W 


24.8.64 


ES 


Chain 50 


76 


2862 


3 


39"38.3'N 


67'57.8'W 


29.6.65 


ES 




77 


3806 


752 


38°00.7'N 


69T6.CW 


30.6.65 


ES 




78 


3828 


199 


38°00.8'N 


69T8.7W 


30.6.65 


ES 




80 


4970 


1 


34°49.8'N 


66"34.0'W 


2.7.65 


ES 




85 


3834 


1 150 3759.2'N 


69"26.2'W 


5.7.65 


ES 


Atlantis II 


124 


4862 


1 


3T26.CN 


63°59.5'W 


22.8.66 


ES 


24 


126 


3806 


48 


39'37.0'N 


66"47.0'W 


24.8.66 


ES 


Atlantis II 


175 


4667- 


1 


36°36.0'N 


68'29.0'W 


28.11.67 


ES 


40 




4693 












Chain 106 


334 


4400 


3 


40°42.6'N 


46'13.8'W 


30.8.72 


ES 




335 


3882- 
3919 


5 


40°25.3'N 


46°30.0'W 


31.8.72 


ES 


Knorr 35 


340 


3264- 95 


38T4.4'N 


70°20.3'W 


24.11.73 


ES 






3356 












BRAZIL BASIN 














Atlantis II 
31 


156 


3459 


6 


00°46.0'S 


2918.0'W 


14.2.67 


ES 


GUYANA BASIN 














Knorr 25 


287 


4980- 
4934 


72 


13T6.0'N 


54"52.2'W 


24.2.72 


ES 




288 


4417- 
4429 


19 


1 1'02.2'N 


55'05.5'W 


25.2.72 


ES 




291 


3859- 
3868 


43 


10°06.1'N 


55"14.0"W 


26.2.72 


ES 




301 


2487- 
2500 


23 


08T2.4'N 


55°50.2'W 


29.2.72 


ES 




303 


2842- 
2853 


8 


08°28.8'N 


56°04.5'W 


1.3.72 


ES 




307 


3862- 
3835 


15 


12"34.4'N 


58°59.3'W 


3.3.72 


ES 


J.Charcot 


DS05 5100 


3 


10"46.0'N 


42°40.3'W 


19.11.77 


DS 


Biovema 
















ARGENTINE BASIN 












Atlantis II 


242 


4382- 


1 


38T6.9'S 


5r56.1'W 


13.3.71 


ES 


60 




4402 














243 


3815- 
3822 


3 


3736.8'S 


52"23.6'W 


14.3.71 


ES 




247 


5208- 
5223 


6 


4333.CS 


48*58. l'W 


17.3.71 


ES 




256 


3906- 
3917 


8 


37'40.9'S 


52T9.3'W 


24.3.71 


ES 




259 


3305- 

3317 


11 


3713.3'S 


52°45.0'W 


26.3.71 


ES 



Distribution. An abyssal species, found widely within temper- 
ate and tropical latitudes at depths ranging from 2487m to 5223m. 



Shell description (Figs 12 & 13) 

Shell robust, ovate, moderately wide, ornamentated with marked 
concentric ridges, straw-coloured periostracum; umbos prominant, 
inwardly facing, clearly anterior of vertical midline in specimens 
>3.0mm, more central in smaller specimens (post-umbonal length 
c. 57% of total length); no clearly marked lunule or escutcheon, 
but some specimens with faint indications; postero-dorsal margin 
slight concave curve, very slightly angulate opposite limit of 
hinge plate then steepening to posterior margin, posterior margin 
may be somewhat flattened particularly in small specimens, ven- 
tral margin moderately curved joining anterior and antero-dorsal 
margins in a smooth curve, anterior and posterior limits of shell 
dorsal to mid-horizontal line; hinge plate broad, continuous, elon- 
gate, short chevron-shaped hinge teeth, up to 14 in posterior 
series, 12 in anterior series, numbers varying with size of speci- 
men, edentulous space below umbo very small; external ligament 
short, opisthodetic, resilium microscopic, close to shell margin, 
ventral to umbo. Maximum total length of present specimens is 
9.8mm. 

Neilonella whoii most closely resembles N. salicensis. It can be 
separated from the latter species by its more rounded shape and 
greater height. The posterior shell margin is not as acute and the 
posterior limit of the shell is more dorsal in position as compared 
with N. salicensis. Furthermore, the post-umbonal length of N. 
whoii is somewhat longer than in N. salicensis and the hinge plate 
is not so broad having a smaller ratio of anterior to posterior 
teeth. 

We name this species in honour of the Woods Hole 
Ocenanographic Institution, through whose auspices these studies 
were carried out. 



Internal morphology (Figs 14 & 15) 

For the most part the morphology of N. whoii is similar to that of 
N. salicensis. Of the mantle structures, the construction of the siphon 
is similar, although the siphonal embayment is less deep than in N. 
salincensis. The posterior adductor muscle is oval in cross-section 
and not much smaller than the anterior, probably reflecting the more 
rounded shell outline ofN. whoii. The anterior sense organ is poorly 
developed, the least developed of all the deep-sea nuculanids that 
have been described to date. The gills and palps of N. whoii are 
similar in size and form to those in N. salicensis with up to a 
maximum of 18 gill plates and 17 palp ridges. 

The foot is large with a few moderately deep papillae at the 
margin. The mouth lies some distance posterior to the anterior 
adductor. The stomach is large with 9 or 10 ridges forming the 
porterior sorting area. The hind gut makes a single loop to the right 
side of the body, the loop being somewhat larger and more smoothly 
curved than that in N. salicensis. The ganglia and commissures are 
not so stout as they are in the latter species. 

Clearly Neilonella whoii is closely related to N. salicensis. 
Although similar in form, they have markedly different depth 
distributions, N. salicensis occurring mainly at lower slope 
depths and N. whoii occurring mainly at abyssal depths. We believe 
that in the past there may have been misidentifications, and speci- 
mens of N. salicensis recorded from abyssal depths deserve 
re-examination. 

Specimens which are narrower and relatively smaller in height to 
length ratio than those described above (Fig. 1 6) are present in some 
samples and do not occur other than with typical specimens of N. 
whoii. In other respects they are no different in their morphology to 
N. whoii. We consider them to be varients at the limit of a range of 
shell outlines and not a subspecies. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



111 







Fig. 12. Neilonella whoii. Details of shell form of specimens, from Chain station 78 from the North America Basin, the type locality; a. left lateral, b, 
dorsal and c, anterior views intact shells; d, the hinge plate of a specimen, from Incal station DS05 from the West European Basin; e, detail of the 
umbonal region of the hinge plate of a specimen, from Chain station 78. Scales = 1mm. 





Fig. 13. Neilonella whoii. Four shells in outline, from Atlantis II station 
72 from the North America Basin, in left lateral view to illustrate the 
small changes in shape with increasing size. Scale = 1mm. 



Fig. 14. Neilonella whoii. A semidiagrammatic view of the internal 
morphology as seen from the right side. For the identification of the 
parts see text-figure 7, p. 106. Scale = 1mm. 



112 





Fig. 15. Neilonella whoii. The dissected stomach and style sac as seen 
from a, in left lateral and b, frontal view. See abbreviations to text 
figures on p. 102. Scale = 1mm. 




f U4<± 



B 




Fig. 16. Neilonella whoii. a, right lateral view of shell of elongate form, 
from Atlantis II station 242 from the Argentine Basin, in right lateral 
view and b, detail of the umbonal region of the same shell in dorsal 
view. Scales = 1 mm. 



J.A. ALLEN AND H.L. SANDERS 

Family Nuculanidae Adams and Adams 1858 

A recent definition of the family is given by Allen, Sanders and 
Hannah (1995). 

Subfamily Ledellinae Allen and Sanders 1982 

The subfamily is defined by Allen and Hannah ( 1 989) and comprises 
two genera, Ledella and Tindariopsis. 

Genus LEDELLA Verrill and Bush 1897 

Type SPECIES. Ledella bushae Waren 1978. SD - Waren 198 1 . 

Shell small, short, robust, surface matt, concentric sculpture, in 
some species scattered incomplete radial striae, usually rostrate, 
single postero-dorsal ridge in some species, postero-ventral margin 
very slightly sinuous, ventral margin in older specimens maybe 
flattened, anterior and posterior hinge teeth series separated by 
edentulous space bearing short internal amphidetic ligament 
(resilium) which may be restricted to dorsal portion of hinge plate, 
outer layer of ligament visible externally and maybe extended 
anteriorly and posteriorly for a short distance, hind gut with various 
configurations. 



Genus TINDARIOPSIS Verrill and Bush 1897 

TYPE SPECIES. Malletia (Tindaria) agatheda Dall 1 889. OD. 

Shell veneriform, matt surface, concentric sculpture, umbo large, 
short rostrum defined by slight radial ridge and furrow, postero- 
ventral margin slightly sinuous, anterior and posterior hinge teeth 
separated by very small edentulous space, internal ligament small, 
close to shell margin, external ligament robust, amphidetic. 

Ledella acinula (Dall 1890) 

Type SPECIMEN. Holotype USNM 95438. 

Type LOCALITY. U.S. Fish Commission Steamer 'Albatross' Sta. 
2754, 1 1"40'N 58°33'W, East of Tobago, 1609m. 

Cited specimens. BMNH 1995047 

Malletia (Tindaria) acinula Dall, 1890, 253, pl.XIII, fig.4. 

Tindaria acinula Verrill and Bush 1898, 881. 

'Tindaria' acinula Sanders and Allen 1977, 55, figs 44, 45. 

Material: 



Cruise 



Sta Depth No Lat 
(m) 



Long 



Date 



Gear 



1 



0758.0'S 34'28.0'W 20.2.67 ES 

08"58.0'N 54"04.3'W 27.2.72 ES 
08'01.9'N 54"16.4"W 28.2.72 ES 



BRAZIL BASIN 
Atlantis II 167 943- 

31 1007 

GUYANA BASIN 
Knorr25 293 1456- 2 
1518 
295 1000- 4 

1022 
299 1942- 108 0755. l'N 55"42.0"W 29.2.72 ES 
2076 
WEST EUROPEAN BASIN 

Chain 106 330 4632 3 50°43.5'N 1751.7'W 24.8.72 ES 
J.Charcot DS79 3226 15 46'30.4'N 10'27.1'W 26.10.74 DS 
Biogas VI 
GUINEA BASIN 

J.Charcot DS28 1261 4 04'21.2'N 04°35.2'E -.-.71 DS 
Walda 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 
The type specimen has been examined by us. 

Distribution. Found predominantly in tropical and subtropical 
latitudes, and southern temperate latitudes in the eastern Atlantic, at 
mid-slope to abyssal depths ranging from 943m to 4632m. 

Many protobranch species with robust shells having subrostrate 
or ovate outlines and with concentric ridges and external liga- 
ments, in the past have been referred to the genera Malletia, 
Neilonella orTindaria (e.g. Dall, 1890). Ledella acinula is a case 
in point. Having addressed this problem in earlier papers (Sand- 
ers and Allen, 1977, 1985), and the present, we have been able to 
define more rigorously the families Tindariidae. Neilonellidae 
and Malletiidae. 

Dall ( 1 890) who described large specimens of L. acinula referred 
them to the subgenus Tindaria. Large specimens do show some 
resemblance to neilonellids and tindariids, but had Dall seen the 
shells of smaller specimens (Fig. 1 8), he would have been unlikely to 
have made the error. 

In our studies on Tindaria (Sanders and Allen, 1977) we re- 
examined and briefly redescribed L. acinula, and recognized that 
there was a problem in identification but deferred final judgment 
until we had made further comparative studies. Now that the 
Ledellinae have been reported upon (Allen and Hannah, 1989), the 
taxonomic relationship of this species is clear. 

ThatL. acinula is siphonate clearly distinguishes it from members 
of the family Tindariidae. Although similar to the neilonellids in 
having an external amphidetic ligament, it differs in having a small 
but well-defined internal ligament and in being semi-rostiate with a 
slightly sinuous postero-ventral margin. A further significant differ- 
ence is the form of the hind gut and the course that it takes within the 
body (Sanders and Allen, 1977). In L. acinula the hind gut is not 
particularly wide in diameter and is not restricted to the right side of 
the body (see below) having a configuration only known to occur in 
species of the subfamily Ledellinae e.g. Ledella galatheae Knudsen 



113 

1970, L. oxira (Dall 1927), L. acuminata (Jeffreys 1870)( Allen and 
Hannah, 1989). 

Shell description (Figs 17 & 18) 

Shell robust, posteriorly angulate, ornamented with concentric ridges 
particularly well-defined on ventral part of shell, straw-coloured 
periostracum; umbos relatively low in profile, inward facing, anterior 
to midline; posterior rostral region characteristically broad and blunt 
when seen in dorsal view; postero-dorsal shell margin almost straight, 
angulate at posterior limit of hinge plate - particularly so in smaller 
specimens, barely so in larger, posterior margin sharply rounded, 
ventral margin deeply curved, postero-ventral margin slightly sinu- 
ous, particularly in smaller specimens, antero-dorsal, anterior and 
antero-ventral margins form a smooth curve; posterior and anterior 
limits of shell at or slightly ventral to mid horizontal axis, ventral limit 
of shell posterior to vertical axis through umbo; hinge plate elongate, 
broad, anterior and posterior tooth series separated by relatively long 
edentulous space, chevron-shaped teeth acutely angled, up to 10 
anterior and 12 posterior teeth depending on size of specimen; 
ligament amphidetic, external parts short, internal resilium small, 
rounded, occupying upper central part of hinge plate below umbo.The 
maximum total length of the present specimens is 6.0mm. 

The shape of the shell changes significantly with growth (Figs 18 
& 19). While the ratio of height to length remains more or less the 
same, the postero-umbonal length increases from 50% to 60% of the 
total length of the shell. With increasing size the postero-dorsal 
margin also becomes less angulate at the posterior limit of the hinge, 
also the postero-ventral margin becomes less sinuous, at most being 
somewhat flattened. 

Internal morphology 

This has been described and illustrated by us in our earlier studies on 
the family Tindariidae (Sanders and Allen, 1977). Only essential 
features relating to the taxonomy need be mentioned. 

Combined siphons are present and there is a well-developed 




Fig. 17. Ledella acinula. a & b, internal and external views of a left valve, from the type locality Albatross station 2754, USNM 95438; c. internal view 
of left valve of specimen, from Atlantis II station 167 from the Brazil Basin; d & e, lateral view of left side and dorsal view of a shell, from Knorr station 
299 from the Guyana Basin. Scale = 1mm. 



114 



J.A. ALLEN AND H.L. SANDERS 




Fig. 18. Ledella acinula. Four shells in outline, from Knorr station 299 from the Guyana Basin, in right lateral view to illustrate changes in shape with 
increasing size. Scale = 1mm. 



feeding aperture ventral to the siphonal embayment. The adductor 
muscles are relatively large, the anterior ovate and the posterior 
more circular in cross section. The anterior sense organ lies far 
anterior, ventral to the anterior adductor. The palps and gills are 
moderate in size with relatively few ridges (up to 15) and plates (up 
to 17) respectively. The foot has a well-defined neck, this is probably 
related to the relatively large height of the shell. There is a large 
'byssal' gland present in the heel of the foot. The hind gut first makes 
a single loop on the right side of the body before passing to the left 
side of the body between the oesophagus and the inner face of the 
anterior adductor muscle. On the left side of the body the hind gut 
forms a double coil. Because of the anterior penetration of the gut to 
the left side, the mouth is displaced some distance posterior to the 
anterior adductor muscle. 



Ledella aberrata (new species) 

Type specimen. Holotype BMNH 1995045; paratypes BMNH 
1995046. 

Type LOCALITY. Chain cruise 60, station 247, Argentine Basin, 
43"33.0'S 48'58.1'W, 5208-5223m 



70-« 



J o-o . 



o 

■05- 



60- 



PL/ 



50- 



1 r 

2 3 

Length (mm) 



Fig. 19. Ledella acinula. The ratios of height to total length (H/L)(open 
circles) and post-umbonal length to total length (PL/L)(closed circles) 
plotted against total length to show changes in shell proportions with 
increasing length. Specimens from Knorr station 299 from the Guyana 
Basin. 



Material: 














Cruise 


Sta Depth 
(m) 


No 


Lat 


Long 


Date 


Gear 


ARGENTINE BASIN 












Atlantis II 


242 4382- 


2 


38"16.9'S 


51"56.1'W 


13.3.71 


ES 


60 


4402 














247 5208- 


34 


43'33.0'S 


48"58.1'W 


17.3.71 


ES 




5223 














252 4435 


4 


38°29.8'S 


52*09. l'W 


22.3.71 


ES 


GUYANA BASIN 












J.Charcot 


KG13 5100 


1 


10'47.6'N 


42°40.4'W 


20.11.77 


KG 


Biovema 














NORTH AMERICA BASIN 










Chain 50 


85 3834 


1 


3759.2'N 


69*26.2'W 


5.7.65 


ES 


WEST EUROPEAN BASIN 










Chain 106 


330 4632 


3 


5043.4'N 


17'51.7'W 


24.8.72 


ES 


J.Charcot 


DS23 4734 


5 


46*32.8'N 


1071.0'W 


1.11.72 


DS 


Polygas 














Biogas II 


DS32 2138 


1 


4T32.2'N 


08"05.3'W 


17.4.73 


DS 


B togas IV 


DS54 4659 


7 


46'31.1'N 


1 0*29.2' W 


21.2.74 


DS 


Cryos 


DS68 4550 


2 


46°26.7'N 


10*23.9'W 


19.6.74 


DS 


Biogas V 














J.Charcot 


DS78 4706 


18 


46"31.2'N 


10°23.8'W 


25.10.74 


DS 


Biogas VI 


DS79 4715 


17 


46"30.4'N 


10*27. l'W 


26.10.74 


DS 




DS80 4720 


3 


46'29.5'N 


10"29.5'W 


27.10.74 


DS 




DS81 4715 


2 


46"28.3'N 


10°24.6'W 


27.10.74 


DS 


INCAL 


CP10 4823 


1 


4875.5'N 


15'10.7'W 


31.7.76 


CP 




DS11 4823 


2 


48'18.8'N 


15"11.5'W 


1.8.76 


DS 




WS034829 


18 


48"19.2'N 


15°23.3'W 


1.8.76 


WS 




CPU 4823 


1 


48"20.4'N 


15'14.6'W 


1.8.76 


CP 




OS02 4829 


1 


48*19.2'N 


15"15.9'W 


2.8.76 


OS 




WS 104354 


1 


4727.3'N 


09"39.9'W 


11.8.76 


WS 


CAPE BASIN 












J.Charcot 


DS05 4560 


2 


33*20.5 'S 


02*34.9'E 


30.12.78 


DS 


Walvis 















\ 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



115 




Post. 



Fig. 20. Ledella aberraia. a, dorsal view of shell; b. dorsal, ventral and anterior view of thickened shell; c. lateral view of the hinge plate of a left valve, all 
from Atlantis II station 247 from the Argentine Basin; d, lateral view of the hinge plate of a right valve, from Chain station 85. Scales = 1mm. 



Distribution. In temperate and tropical basins of the Atlantic at 
abyssal depths >4000m, rare in the North America Basin. Depth 
range, 2138-5223m. 

Shell description (Figs 20 & 21) 

Shell small, ovate, relatively high, moderately wide, ornamented 
with concentric ridges; umbo moderate in size, inwardly turned, 
anterior to mid-line but less so in juveniles; no lunule or escutcheon; 
periostracum pale straw colour; postero-dorsal margin slightly con- 
vex becoming more straight with increasing size, slightly angulate at 
posterior limit of hinge plate and at posterior margin, postero-ventral 
margin very slightly sinuous, otherwise ventral margin deeply curved 
with ventral limit posterior to vertical axis through umbo, anterior 
margin sharply curved, antero-dorsal margin slightly convex with 
slight change in slope at anterior limit to hinge plate; shell outline 
characteristically asymmetrical, shell margin in larger specimens 
changes direction of growth producing a broad flattened ventral 
margin; hinge plate broad, up to 6 chevron teeth in anterior series and 
7 in posterior series, edentulous space between series relatively 
broad; ligament small, amphidetic, internal part restricted to upper 
part of hinge plate, external part extremely short situated below 
umbo. The maximum length of the present specimens is 2.6mm. 



Internal morphology (Figs 22 & 23) 

The adductor muscles are moderately large and oval in shape. The 
combined siphon is relatively short The dorsal margins of the 
exhalent part are fused proximally for a short distance and the 
ventral margins of the inhalent part are not fused but are slightly 
thickened and probably adhere in living specimens. Internally where 
inhalent and exhalent parts join, there is a thickened median ridge on 
each side which together together with the posterior continuation of 
the gill axes probably act as guides when the faecal pellets are 
extruded. The siphonal embayment is small and there is a small, 
slender, tentacle attached to the left side at the inner limit of the 
embayment. The anterior sense organ is small and is situated ventral 
to the anterior adductor muscle. 

The palps are small with up to 11 broad ridges. The gills are 
also small each with up to 11 plates the most posterior of which 
lies some distance from the siphon. The gills are attached to the 
posterior limits of the median guides by slender extensions of the 
gill axis. 

The hind gut is greatly extended and takes a similar but more 
complex course to that described for L. acinula. Like the latter, the 
hind gut passes from the right side of the body to the left immedi- 
ately posterior to the anterior adductor muscle and returns by the 



116 



J.A. ALLEN AND H.L. SANDERS 




Fig. 21. Ledella aberrata. Five shells in outline to show variation in 
shape with growth, a, is a specimen with a thickened margin; b. has a 
slightly thickened margin; the remaining three shells are unthickened. 
Scale = 1mm. 

same route. Unlike the latter species, it makes a single coil on the 
right side as well as a double twinned coil on the left. 

The foot is unusual in having a large heel and a narrow muscular 
anterior part. The marginal papillae are few in number and restricted 
to the anterior margins of the sole. The sole is less deeply divided as 
compared with other protobranchs. There is a large 'byssal' gland in 
the heel of the foot. 

Although the shell outline of the smaller specimens is more 
characteristic of the genus Ledella than in larger mature specimens, 
the general shell outline of L aberrata (and L. acinula) is much 
deeper and more ovate than in other described species (Figs 1 8 & 2 1 ) 
nor is it markedly rostrate. Despite this, the characters place them in 
the Ledellinae (Allen and Hannah, 1989) and we see no reason for 
erecting another genus. 

Apart from L. aberrata, only two species of protobranchs, have 
been reported as exhibiting a change in shell growth to produce a 
flattened shell margin (Fig. 20). Both are ledellids, namely L. ultima 
(Smith 1885) and L. solidula (Smith 1885) (Allen and Hannah, 
1989). Like L. aberrata these two latter species also have elongate 
hind guts. The hind gut of L. solidula is very similar to that of L. 
aberrata in having double twinned coils to the left side of the body 
(Fig. 22), although it does not have an additional single coil to the 
right as does the present species. The type of course taken by the 
hind gut in L. acinula is also found in other species of Ledella (e.g. 
L. oxira) (Allen and Hannah, 1989). 

We named this species after the familiar appelation to which it 
was referred during our original analysis of the samples. 

Tindariopsis agatheda (Dall 1889) 

Type specimen. USNM 95437, lectotype here designated. 

Type locality. U.S. Fish Commission Sta. 2754, east of Tobago, 
1 1°40'N 58"33W, 1609m. 




Fig. 22. Ledella aberrata. Internal morphology as seen in a, right lateral 
view, b, left lateral view and c, left ventro-lateral view. For identification 
of the parts see text-figure 7, p. 106. Scales = 1mm. 

Cited specimen. BMNH 1995062. 

Malletia (Tindaria) agatheda Dall 1890, 252, pi. xiii, fig. 10. 
Tindaria (Tindariopsis) agatheda Verrill and Bush 1897, 59. 
Saturnia (Tindariopsis) agatheda McAlester 1969, N235. 
Tindariopsis agatheda James 1972, 98, figs 60-62. 
Neilonella (Tindariopsis) agatheda Laghi 1986, 190, pi. 8, figs 2-6. 

Material: 



Cruise 



Sta Depth No Lat 
(m) 



Long 



Date Gear 



BRAZIL BASIN 

Atlantis II 167 943- 1 07'58.0'S 34'28.0'W 20.2.67 ES 

31 1007 

GUYANA BASIN 

Knorr25 293 1456- 14 08"53.1'N 54'04.3'W 27.2.72 ES 
1518 
299 1942- 8 0T55.1'N 55°42.0'W 29.2.72 ES 

2067 
301 2487- 7 08"12.4'N 55'50.2'W 1.3.72 ES 
2500 



The type specimen has been examined by us. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



117 




Fig. 23. Ledella aberrata. Siphonal region, as seen from the inside of the 
mantle cavity. See abbreviations to the text-figures p. 102. Scale = 0.1mm. 

This species occurs at upper to mid slope depths in the tropical 
western Atlantic in the Brazil, Guyana, Caribbean and Gulf of 
Mexico Basins (Dall, 1889; James 1972). Depth range; 943-2500m. 

The holotype for T. agatheda was not designated by Dall ( 1 890), 
nevertheless he did illustrate the left valve from Albatross Sta. 2754, 
1 1°40'N 58"33'W (USNM 95437) which we have here nominated as 



lectotype. We have redrawn the shell and added detail of the hinge 
plate (Figs 24 and 25). In addition, James (1972) reported that two 
valves of T. agathedawere included in USNM 63 149 from Blake sta. 
236, 2909m off Bequia, furthermore USNM 94326 from Blake stas 
26 and 30 between Cuba and Yucatan, identified by Dall as Leda 
pusio, are examples of T. agatheda. 



Shell description (Figs 24 & 25) 

Shell small, sub-ovate, wide, sub-rostrate, ornamented with con- 
centric ridges, postero-lateral furrow ventral to sub-rostrum, 
ill-defined lunule outlined by obscure ridge, escutcheon similarly 
ill-defined and bounded by faint ridge; umbo prominant, poste- 
rior to mid-line, inwardly directed; antero-dorsal margin concave, 
with change in slope at anterior limit of hinge plate, slightly 
flattened anteriorly dorsal to the anterior limit of shell, antero- 
ventral and ventral margins smoothly curved, postero-ventral 
margin slightly sinuate, posterior margin acutely angled, postero- 
dorsal margin slightly concave, marked angle at posterior limit of 
hinge plate and thereafter almost straight to form sub-rostrate 
posterior margin; anterior and posterior limits of shell are ventral 
to horizontal mid-line; hinge plate strong, with edentulous space 
ventral to umbo, hinge teeth stout, chevron-shaped, up to 12 in 
each series in shell 6mm total length; internal ligament small, 
close to shell margin, external ligament amphidetic, moderately 
short, stout. 

Young shells are less rostrate, with the posterior and anterior 
limits of the shell more dorsal in position. 

Although there is variation in the shape of the shell, the height/ 
length and the post-umbonal length/total length ratios increase 
slightly with increasing size (Table 1 ). The maximum total length of 
the present specimens is 6.0mm. 




Fig. 24. Tindariopsis agatheda. External lateral view of the right valve of the lectotype and an internal view of the hinge plate of the same valve, from 
U.S. Fish Commission station 2754 East of Tobago. USNM 95437. Scales = 1mm. 



118 



J.A. ALLEN AND H.L. SANDERS 






Fig. 25. Tindariopsis agatheda. External lateral views of right side of two specimens of differing size to show change in shape with growth, a, from Knorr 
station 301 and b, from Knorr station 293, both from the Guyana Basin: c. external dorsal view of a shell also from station 293. Scale = 1mm. 



Table 1. Measurements and ratios of shell parameters of the sample from 
Knorr sta. 293. 



Length (L) 


Height (H) 


Post-umbonal (PL) 


PL/L 


H/L 


(mm) 


(mm) 


length (mm) 






6.00 


4.95 


3.00 


0.50 


0.83 


5.60 


3.95 


2.65 


0.47 


0.71 


5.00 


3.65 


2.20 


0.44 


0.73 


4.90 


3.65 


2.05 


0.42 


0.75 


4.70 


3.15 


1.95 


0.42 


0.67 


4.50 


3.10 


1.85 


0.41 


0.69 


4.45 


3.20 


1.85 


0.42 


0.72 


3.70 


2.55 


1.70 


0.46 


0.69 


3.45 


2.50 


1.45 


0.42 


0.73 


2.35 


1.58 


0.93 


0.39 


0.67 


2.05 


1.43 


0.93 


0.45 


0.70 


1.95 


1.30 


0.88 


0.45 


0.67 


1.83 


1.28 


0.83 


0.45 


0.70 


1.10 


0.88 


0.48 


0.43 


0.80 



Internal morphology (Fig.26) 

The siphonal embayment is shallow and dorso-ventrally narrow. In 
contrast, the feeding aperture is broad and well-supplied with radial 
pallial muscles. The adductor muscles are small, the posterior 
muscle is oval and the anterior muscle is circular in cross section. 
The foot is large with a well-developed heel containing a large 
'byssal' gland. The gland opens into the posterior limit of the pedal 
groove via a small papilla. The anterior two-thirds of the margins of 
the divided sole are broadly papillate. The palps are large with up to 
25 ridges in the largest specimens. The gills are narrow ill-defined 
with about 16 plates in the largest specimens. 

The mouth lies close to the posterior face of the anterior adduc- 
tor muscle. The oesophagus opens into a large stomach and style 
sac. The hind gut passes posterior to the style sac and stomach to 
the dorsal side of the viscera and thence across the right side of 
the body where it forms 8 coils before returning along the same 
path to the dorsal side of the viscera and from there through the 
heart and then dorsal to the posterior adductor muscle to the anus. 



Usually not more than six coils are visible, the others being 
overlain by those to the outside of them. The form of the hind gut 
is derived by 4 complete turns of the closely parallel anterior and 
posterior lengths of the hind gut on the right of the body. This 
particular disposition of the hind-gut is to be found in other 
ledellids (e.g. L. ultima) and yoldiellids (e.g. Y. ellaAWen, Sanders 
and Hannah 1995) (Allen and Hannah, 1989; Allen, 1992; Allen, 
Sanders and Hannah, 1995). 

The nervous system is similar in its arrangement to that of other 
deep-sea protobranchs, however the cerebral and visceral ganglia 
are noticably smaller and the commissures much finer than observed 
in other species. 




Fig. 26. Tindariopsis agatheda. Internal morphology as seen from the 
right side of a specimen from Knorr station 293. For identification of the 
parts see text-figure 7. p. 106. Scale = 1mm. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 

Tindariopsis aeolata (Dall 1890) 

Type specimen. Holotype, USNM 95436. 

Type locality. U.S. Fish Commission Sta. 2754, East ofTobago, 
1 1'40'N 58"33'W, 1609m. 

Cited specimen. BMNH 1 99506 1 . 

Malletia (Tindaria?) aeolata Dall 1890, 252. 
Tindaria (Tindariopsis) aeolata Dall 1898, 582. 
Tindariopsis aeolata James 1972, 97, figs 57-59. 

Material: 



Cruise 



Sta 



Depth No Lat 
(m) 



Long 



Date 



Gear 



GUYANA BASIN 

Knorr25 299 1942-4 07"55.I'N 55"42.0'W 29.2.72 ES 
2076 
301 2487-5 08'12.4'N 55"50.2'W 29.2.72 ES 
2500 



The type specimen has been examined by us. 

This species occurs from mid to lower slope depths in the tropical 
western Atlantic in the Guyana and Caribbean Basins and the Gulf of 
Mexico. Depth range: 1609-3466m. 

Shell description (Figs 27 & 28) 

Shell small, subquadrate, rostrate, ornamented with marked concen- 
tric ridges; periostracum pale yellow; umbos moderately large, 
posterior to midline (post-umbonal length 45-48% of total length), 
facing inwards, slightly separated by external ligament; distally 
antero-dorsal shell margin horizontal, then curves smoothly and 
steeply to anterior margin, postero-ventral margin sinuous, ventral 
margin somewhat flattened, postero-dorsal distal margin slopes 
gently to limit of hinge plate then curves sharply to rostral point, 
latter rounded and somewhat eroded in large specimens, more 
pointed in smaller, rostrum in mid horizontal plane in small speci- 
mens and dorsal to it in large specimens, limit of anterior margin 





Fig. 27. Tindariopsis aeolata. External lateral view of the right valve and 
the hinge plate of the left valve of the holotype, from U.S. Fish 
Commission station 2754. USNM 95436. Scale = 1mm. 






Fig. 28. Tindariopsis aeolata. External lateral views of the right side of 
shells of differing size to change in shape with growth, a, from Knorr 
station 301 and b, from Knorr station 299 from the Guyana Basin; c, 
external dorsal view of a shell also from Knorr station 299. Scales 
= 1 mm. 

ventral to the mid horizontal plane; hinge plate stout, small edentu- 
lous space between tooth series, 9 chevron-shaped teeth in anterior 
series and 10 in posterior series in largest specimens; ligament 
amphidetic, external except for small resilifer at margin ventral to 
umbo, external part thickened, particularly so in large specimens. 
The maximum length of the present specimens is 5.8mm. 

In lateral view the rostrum, although dorsal to mid horizontal line, 
is reminiscent of Ledella, while the robust external ligament is more 
reminiscent of Spinula. 

Internal morphology (Fig.29) 

The siphonal embayment is relatively shallow and the contained 
siphon is similar to that of T. acinula. The adductor muscles are 
moderately small, ovate in cross-section and equal in size. The foot 
is large, with a divided sole fringed with large papillae. The 'byssaF 
gland is moderate in size. The palps are very large with many ridges 
(c 26 in the largest specimen) and the palp proboscides are broad. 
The gills are small with 1 1 gill plates in the largest specimen. The 
kidney is long and narrow. The nervous system is of typical 
protobranch design. The ganglia are relatively large and, in contrast, 
the commissures are unusually slender. 

The mouth lies some distance posterior to the anterior adductor 
muscle (see below). The oesophagus opens on to the anterior face of 
a moderately large stomach. The latter lies almost vertical within the 
posterior part of the visceral mass. The hind gut is very small in 
diameter and takes an extraordinarily complex course through the 
body. There are two loops to the left side of the body (Fig.29B) and 
one major loop to the right side of the body, all three pass from one 
side to the other ventral to the umbo. There is also a complex series 
of loops anterior and to the right of the stomach. This morphology 
has not been encountered before in the protobranch bivalves and is 
very different from that seen in T agatheda. Yet, it is debateable 



120 



J.A. ALLEN AND H.L. SANDERS 




Fig. 29. Tindahopsis aeolata. a, internal morphology as seen from the 
right side of a specimen from Knorr station 301 from the Guyana Basin; 
b, the form of the hind gut on the left side of the body. For identification 
of the parts see text-figure 8, p.. Scale = 1mm. 

whether the difference warrents generic status. Other protobranch 
genera show an array of hind gut morphologies (e.g. Yoldiellidae, 
Allen, 1992; Allen, Sanders and Hannah, 1995) which we believe 
relate to changes in the benthic food resource as depth increases. For 
this reason we are reluctant to erect a new genus when in other 
respects T. aeolata is clearly within the genus Tindariopsis. 



Subfamily Nuculaninae Allen and Sanders 1982 

The subfamily is defined by Allen and Hannah ( 1 986) and comprises 
three genera Nuculana, Propeleda and Adrana. 



Genus NUCULANA Link 1807 

TYPE SPECIES. Area rostrata Gmelin 1791 = Area pernula Miiller 
1779. OD. 

Shell robust, moderately elongate, concentric sculpture, occa- 
sionally with radial ribs, slightly rostrate, usually bicarinate; umbo 
anterior; postero-dorsal margin straight or somewhat concave, pos- 
terior margin may be slightly sinuous; escutcheon present; no 
internal ridge from umbo to posterior margin; hinge moderately 
robust, teeth chevron-shaped; ligament small, for most part internal, 
usually amphidetic and vertical, sometimes posteriorly oblique. 



Genus PROPELEDA Iredale 1924 

TYPE SPECIES. Leda ensicula Angas 1877. OD. 

Shell very elongate, thin, glossy, concentric sculpture may be ill- 
defined, 3/4 shell post-umbonal, usually with two marked carinae 
from umbo to upper and lower limit of rostrum, posteriorly truncate; 
umbo small; postero-dorsal margin concave, postero-ventral margin 
not sinuous; internal ridge usually from umbo, skirts ventral margin 
of posterior adductor to posterior margin, second ridge may be 
present from hinge plate to rostral margin; hinge plate slender, hinge 



teeth chevron-shaped, one or both arms of the chevron may be 
elongate, anterior tooth series curve round the outer margin of the 
anterior adductor, posterior series extends posterior to adductor; 
ligament in large part internal, opisthodetic and oblique. 



Genus ADRANA Adams & Adams 1 858 

Type SPECIES. Nucula lanceolata Lamarck 1819. SD Stoliczka 
1871. 

Shell extremely elongate, slender, lanceolate, fragile, smooth or 
with fine concentric and sometimes oblique sculpture, without 
carinae, glossy; umbo almost central, barely raised; escutcheon 
elongate, flattened, narrow; postero-dorsal margin straight, antero- 
dorsal margin slightly convex, postero-ventral margin sinuous; hinge 
plate slender, hinge teeth fine, obtuse, chevron-shaped ;chondrophore 
present; ligament internal, amphidetic. 

Nuculana acuta (Conrad 1831) 

Type specimen. Lectotype here designated, chosen from ANSP 
30613, remainder of lot designated paralectotypes. 

Cited specimen. BMNH 1995055. 

Type locality. Tertiary fossil beds, near Suffolk, Virginia. 

Nucula acuta Conrad 1 83 1 , 32, pi. 6, fig. 1 . 
Nucula cuneata Sowerby 1833, 198. 
Nucula carinata H.C.Lea 1843, 163, (non M'Coy 1844). 
?Ledajamaicensis d'Orbigny 1846, 263, pl.XXIV, figs 30-32. 
?Leda inornata A.Adams 1856, 48. 
Leda unca Verrill 1880, 401, (?non Gould 1862). 
Leda acuta Dall 1886, 251, pl.7, figs 3a, 3b and 8. 
Nuculana acuta Morris 1951, 7, pi. 6, fig. 2. 

Material: 



Cruise 



Sta 



Depth No Lat 
(m) 



Long 



Date 



Gear 



NORTH AMERICA BASIN 

Atlantis CI 97 2+2v 40°20.5'N 70'47.0'W 25.5.61 AD 

264 

Atlantis Slope 200 8+6v 40*01. 8'N 70"42.0'W 28.8.62 AD 

283 sta.2 

Atlantis II 114 197 8 40*04. l'N 70"27.8'W 15.8.66 ES 

24 

Atlantis II 172 119 7+2v 40'12.3'N 70"44.7'W 27.1 1.67 ES 

40 173 123 4 40°10.8'N 70"43.6'W 28.1 1.67 ES 



The type specimens have been examined by JAA. 

Conrad (1831) described this species from fossils obtained from 
the Miocene beds near Suffolk and the banks of the James and York 
rivers, Virginia. He later redescribed the species (Conrad, 1845) 
adding that he had found Recent specimens in deep water in the Gulf 
of Mexico. His first account refers to specimens being in 'Cabinet of 
the Acad. Nat. Sciences, No. 1738.' This reference does not corre- 
spond with any lot of N. acuta, Recent or fossil, in the Academy of 
Natural Sciences, Philadelphia today. There are specimens in the 
Invertebrate Paleontology collection of the Academy (catalogue 
number 30613) that up to now have been considered as possible 
syntypes of the species. The lot comprises 4 right valves, 3 left 
valves 1 left and 1 right broken valve, 1 intact shell, 1 shell with 
rostrum tip missing and 3 fragments. These specimens, labelled by 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



121 




Fig. 30. Nuculana acuta, a & b, lateral and dorsal external views of the lectotype, ANSP 3061 3; c, lateral internal view of a paralectotype from the same 
lot. Scales = 1mm. 



Conrad, may include those figured by him. Comparison with the 
figures (Conrad, 1831 and 1845) shows that it is impossible to say 
which, if any one, was figured nor is it possible to be absolutely 
certain that these are the specimens from which the original descrip- 
tion was made, although we believe that they are. Thus, the intact 
shell from lot 3061 3 has been chosen as the lectotype, the remainder 
being paralectotypes. 

Campbell (1993) listed Leda jamaicensis d'Orbigny 1846, Lecla 





Fig. 31. Nuculana acuta. Lateral and dorsal external views of a shell, 
from Atlantis 283 station 2 from the North America Basin. Scales = 
lmm. 



inornata A.Adams 1 856 and L.unca Gould 1 862 as synonyms of A 7 . 
acuta. Verrill (1882) describes in detail differences that he found 
between N. acuta and L.jamaicensis and L. unca which cast doubt as 
to the synonymy, although Dall (1886) maintains the synonymy of 
L. jamaicensis. Similarly, we have doubts as to synonymy with 
L.inornata A.Adams which is a 'gibbose', 'fuscous', 'sulcate' spe- 
cies from New Guinea, Thus, although Nuculana acuta is a 
well-described species (e.g. Verrill 1882, 1884; Dall, 1886; Abbott, 
1974) because there are closely related species in the Atlantic and 
elsewhere, we include a description here. It occurs off the East coast 
of North America, in the Caribbean Sea and off Brazil at depths from 
the outer shelf to lower slope depths, 97-2909m (James, 1972). 

Shell description (Figs 30, 31 ) 

Shell moderately large, elongate, somewhat inflated, rostrate, 
ornamented with deep concentric ridges flattened at the apex, rostral 
ridge from umbo to ventral limit of rostrum, very faint radial ridge 
from umbo to antero-ventral margin, yellow periostracum; lunule 




Fig. 32. Nuculana acuta. Internal morphology as seen from the right side 
of a specimen from Atlantis II station 197 from the North America Basin. 
For identification of the parts see text-figure 7, p. 106. Scale = lmm. 



122 



J.A. ALLEN AND H.L. SANDERS 




D 




Fig. 33. Nuculana acuta. Internal morphology, a, anterior, b, left lateral & c. posterior views of a dissected stomach and combined style sac; d, ventral 
view of siphonal region. See abbreviations to text-figures on p. 102. Scales = 1mm. 



broad, elongate, outlined with faint ridge, escutcheon broad, out- 
lined by rostral ridge; umbos small, inwardly directed, anterior to 
mid-line; antero-dorsal margin broadly concave, anterior, antero- 
ventral, ventral and postero-ventral margins form smooth curve, 
postero-dorsal margin raised, distally straight, proximally - poste- 
rior to hinge plate - slightly concave, may be slightly upturned in 
larger specimens; hinge plate elongate, relatively broad, hinge teeth 
chevron-shaped, up to 18 teeth in each series depending on size of 
specimen; ligament small, amphidetic, internal pear-shaped in sag- 
ittal section, extends slightly ventral to hinge plate. The maximum 
length of the present specimens is 9.2mm. 

Internal morphology (Figs 32, 33) 

The siphonal embayment is deep, with an elongate tentacle attached 
to the inner right or left side. The siphons are elongate, combined 
and except anteriorly, the ventral margins are fused. In the con- 
tracted state the line of fusion is marked by deep ventral furrow. The 
anterior sense organ is far anterior, situated at the point where the 
radial ridge meets with the shell margin. Between the feeding 
(ventral to the siphonal embayment) and the pedal gape, the inner 



folds of the ventral margin are applied to each other. In this section 
of mantle margin approximately 30 small sensory papillae are 
attached to each middle sensory fold in a specimen 6.5mm total 
length. The adductor muscles are small the anterior unusually so. 
The anterior adductor is circular in cross-section and the posterior 
elongate-oval. 

The foot is moderately large, elongate, the sole with papillate 
margins. The heel is not marked but there is a a large 'byssal' gland 
present internal to the posterior limit of the sole. The gills are well- 
developed with up to 48 alternating gill plates. The dorsal margins of 
the left and right inner demibranchs are fused. In life the gills are a 
bright orange-red colour. In contrast the palps are cream. The latter 
are relatively small, elongate and dorso-ventrally narrow and. for the 
most part, hidden under a fold of the body wall. This latter is more 
pronounced on the right side of the body where the hind gut loop 
meets the ventral margin of the visceral wall. Each palp has up to 24 
ridges on the inner face. The palp proboscides are also long and 
tapering. 

The digestive gland is bright orange in life. As in other species of 
Nuculana, the course of the hind gut describes a single loop to the 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



123 



right side of the viscera. There is a single typhlosole present along its 
entire length. The stomach is of moderate size and internally is 
similar in form to that of shallow water species of Nuculana ( Yonge, 
1939). The gastric shield lines much of the left wall of the stomach. 
To the right there is a large posterior sorting area with 13 ciliated 
ridges. A deep caecum is ventral to the oesophageal aperture. Two 
digestive ducts open close to the antero-dorsal margin of the poste- 
rior sorting area and a single duct opens antero-dorsally close to the 
oesophageal opening. 

As will be seen Nuculana acuta is remarkably similar in its shell 
features and anatomy to Nuculana commutata. This similarity is 
discussed under the latter species (p. 123). 

Nuculana commutata (Philippi 1844) 

Type specimen. ZMHU. 

Type LOCALITY. Pliocene, Palermo, Sicily. 

Cited specimen. BMNH 1995212 

Area fragilis Chemnitz 1784, 199, pl.LV, fig.546. 
Area pellet Gmelin 1790 (non Linne), 3307. 
Area minuta Brocchi 1814 (non Fabricius), 482, pl.XI,fig.4. 
Nueula pella Payraudeau 1826 (non Linne), 64. 
Lembulus eieltoieleus Risso 1826 (non Lamarck), 320, pi. XI, fig. 

164. 
Nueula minuta Scacchi 1836 (non Fabricius), 4. 
Nueula striata Philippi 1836 (non Lamarck), 64. 
Nueula commutata Philippi 1844, 101. 
Leda fragilis Jeffreys 1879, 575. 
Leela minuta Jeffreys 1856 (non Fabricius), 25. 
Leda commutata Hanley 1863, pl.CCXVIII, figs, 80, 81. 
Lembulus commuteitus Monterosato 1878, 6. 
Leela (Portlandia) tenuis Sturany 1 896, 6. 
Nuculana (Jupiteria) fragilis Nordsieck 1969, 9, pi. I, fig. 02. 25. 
Nuculana (Jupiteria) commutata Smith and Heppell 1991, 56. 



Material: 








Cruise Sta Depth No 

(m) 


Lat 


Long 


Date Gear 


WEST EUROPEAN BASIN 
Sarsia S29 119 16 


4740.0'N 


05"00.0'W 


12.8.67 ES 



Shell description (Fig. 34) 

Shell moderately large, elongate, slightly inflated, rostrate, 
ornamented with concentric ridges, pale straw-coloured perio- 
stracum; radial ridge from umbo to antero-ventral margin; rostral 
ridge well-defined, delimits posterior dorsal area, within this area a 
faintly outlined escutcheon extending half the length of postero- 
dorsal margin; lunule elongate, defined by fine ridge; less 
well-defined ventral rostral ridge extends from umbo to postero- 
ventral margin; umbos anterior to midline, inwardly directed; 
antero-dorsal margin proximally straight, distally slightly concave 
merging with rounded anterior margin to where it meets with ventral 
limit of anterior radial ridge, ventral margin broadly concave, postero- 
ventral margin sinuate where ventral rostral ridge meets margin, 
posterior margin acute, slightly upturned, postero-dorsal margin 
slightly raised with shallow angulation at limit of hinge plate; hinge 
plate elongate, relatively broad, acute chevron teeth, 16 on both 
anterior and posterior hinge plates of specimen 8.3mm total length; 
ligament internal, amphidetic, triangular, extends slightly ventral to 
hinge plate. 




Fig. 34. Nuculana commutata. Lateral and dorsal views of a shell, from 
Sarsia station S29 from the West European Basin. Scale = 1mm. 
Maximum length of present specimens is 8.3mm. 

Internal morphology (Fig. 35) 

The internal morphology differs little from that ofN. acuta (Fig. 32). 
The most noticable differences are that N. commutata has less 
attenuate palps with fewer palp ridges and larger adductor muscles 
than does N. acuta. 

Other differences between the two species are that inN. commuteita 
the ridge from the umbo to the antero-ventral margin is more 
marked, the apices of the concentric ridges are less broad, the 
postero-dorsal margin is not so raised and the shell is somewhat less 
elongate in relation to its height. 

These differences are of degree and at that point where separation 
into species rather than subspecies is a subjective judgement. Never- 
theless, these differences are more marked than those between N. 
commutata andN. ////WcaCarrozza 1987 (paratypes BMNH 1995213 
examined by JAA), a species that has been recently described from 
the northern Adriatic Sea (Carrozza, 1987). In contrast^ commutata 




Fig. 35. Nuculana commutata. Internal morphology as seen from the 
right side of a specimen from Sarsia station S29 from the West 
European Basin. For identification of the parts see text-figure 7, p. 106. 
Scale = 1mm. 



124 



J. A. ALLEN AND H.L. SANDERS 



is widely distributed throughout the Mediterranean and lusitanean 
Atlantic. Prior to the paper by Carrozza ( 1 987) there had been debate 
as to whether N. fragilis and N. commutata were the same species 
(Locard, 1891, 1898; Bucquoy et al, 1887-98). It is not possible to 
determine whether this earlier debate was a presage to the study of 
Carrozza (1987). In contrast N. acuta is even more widely distrib- 
uted off the eastern North America, West Indies and off Brazil 
(Abbott, 1974). It must be assumed that these are three sibling 
species. 

Nuculana vestita (Locard 1898) 

Type specimen. MNHN 

Type LOCALITY. Talisman stas 96-98 & 101, West of Senegal, 
2324-3200m, 19'12'N 1757'W - 16°38'N 18°24'W 

Cited specimens. BMNH 1 995056 and 1 9952 1 1 

Leda vestita Locard 1898, 340, pl.XIV, figs 12-18 

Nuculana vestita Clarke 1962, 53. 

Leda macella Barnard 1963, 448, fig. lid; type locality: West off 

Cape Point, S.W.Africa, Africana II stas A 190, A 192, A3 17, 

A3 19, 2268-3200m, SAM (not seen). 



MATERIAL: 












Cruise Sta Depth 
(m) 


No 


Lat 


Long 


Date 


Gear 


SIERRA LEONE BASIN 












Atlantis II 146 2842 


1 


10"39.5'N 


17°44.5'W 


6.2.67 


ES 


31 -2891 












ANGOLA BASIN 












Atlantis II 201 1964 


1 


09"29.0'S 


1 1°34.0'E 


23.5.68 


ES 


42 -2031 












203 527 


742 


0846.0'S 


12'47.0'E 


23.5.68 


ES 


-542 













Specimens taken by the Galathea Expedition described by 
Knudsen (1970) examined by JAA, ZMUC. Knudsen (1970) fol- 
lowing examination of specimens synonymized L. macella with N. 
vestita. 

Nuculana vestitais a well-described species (Locard, 1 898;Theile 
and Jaeckel, 1931; Knudsen, 1970) occurring off West and South- 
west Africa at lower slope depths (71 5-289 lm) in the Sierra Leone, 
Guinea and Angola basins. 



Shell description (Figs 36 & 37) 

Shell moderately large, inflated, somewhat elongate, rostrate, 
ornamented with concentric ridges, pale brownish-yellow 
periostracum; umbos large, inwardly directed, anterior to midline; 
antero-dorsal distal margin horizontal for short distance, proximal 
margin broadly convex forming smooth curve with anterior margin, 
ventral margin broadly convex to rostrum, postero-dorsal margin, 
raised in small specimens but less so in large, proximally straight or 
slightly concave, in small specimens angulate at limit of posterior 
hinge plate, distally slightly concave to posterior limit of rostrum; 
broad ridge extends from umbo to rostrum forming outer limit of 
escutcheon; anterior and posterior hinge plates broad, meet shell 
margin ventral to umbo, hinge teeth broad chevrons, up to 19 and 16 
in anterior and posterior series respectively in specimen 8.3mm total 
length; ligament internal, amphidetic, pear-shaped in sagittal verti- 
cal section, extends ventral to hinge plate. The maximum length of 
the present specimens is 13.8mm. 

Internal morphology (Fig. 38) 

The adductor muscles are moderately large and oval. The siphonal 
embayment is deep with the siphonal tentacle to the right side. The 
siphons are entire. The feeding aperture is small but well-defined 
with the mantle surface ridged internally. Radial mantle muscles 
are well-developed forming a broad band internal to the inner lobe 




Fig. 36. Nuculana vestita. External right lateral and dorsal view of a large adult shell and a lateral view of the hinge plate of a right valve. Specimens are 
from Atlantis II station 203 from the Angola Basin. Scale = 1 mm. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



125 




Fig. 37. Nuculana vestita. Lateral views of right side of four shells in outline to show differences in shape with increasing size. Specimens from Atlantis 
II station 203 from the Angola Basin. Scale = 1mm. 



of the mantle edge. The anterior mantle sense organ is well- 
developed. 

The foot is large with the margins of the sole finely papillate. The 
palps are very small with up to 12 broad folds. The palp proboscides 
are stout and elongate, even in the contracted state. The gills are 
elongate and broad with up to 36 gill plates. 

Propeleda carpenteri (Dall 1881) 

Type specimen. Syntypes, USNM 63 1 5 1 and MCZ 7936-7938. 

Type locality. Off Barbados, 'Blake' stations 5, 9, and 21, 
100fm-287fm. 

Cited specimens. BMNH 1995057 

Leda carpenteri Dall 1881, 125; 1886, 249, pi. 8, fig. 10, pl.9, fig.3. 
Nuculana carpenteri Johnson 1934, 16. 

Material: 



Cruise 



Sta Depth No Lat 

(m) 



Long 



Date Gear 



ARGENTINE BASIN 

Atlantis II 237 993- 194 36"32.6'S 53'23.0'W 11.3.71 ES 
60 1011 

239 1661- 8 36'49.0'S 53"15.4'W 11.3.71 ES 
1679 

240 2195- 8 36°53.4'S 53"10.2'W 12.3.71 ES 
2323 



Specimen USNM 63151 examined by JAA. 

Previously reported off N. Carolina, Gulf of Mexico and Eastern 
Caribbean (Dall, 1889; Rice and Kornicker, 1965; James, 1970), the 
present specimens are from the Argentine Basin. This species has a 




Fig. 38. Nuculana vestita. Internal morphology as seen from the right 
side of a specimen from Atlantis II station 203 from the Angola Basin. 
For identification of the parts see text-figure 7, p. 106. Scale = 1mm. 

somewhat unusual distribution from shelf to lower slope depths, 
200-2323m 

Shell description (Figs 39^-1) 

Shell fragile, semi-transparent, slender, moderately elongate, rostrate 
with two post umbonal carinae, ornamented with faint concentric 
ridges; periostracum pale straw colour; umbos small, far anterior 
(postumbonal length 60-68% of total length), inwardly facing; 
antero-dorsal margin slightly flattened, ventral margin smoothly 
curved, postero-dorsal margin raised, slightly sinuous, concave 
proximally, convex distally, meets posterior margin at limit of dorsal 
post-umbonal carina, posterior margin concave between posterior 



126 



J.A. ALLEN AND H.L. SANDERS 




Fig. 39. Propeleda carpenteri. External lateral view of the left valve and an internal view of the same valve of a syntype USNM 63151, from off 
Barbados in lOOfms. Scale = 1mm. 



limits of dorsal and ventral carinae; escutcheon lanceolate; hinge 
plates relatively broad, posterior plate short, occupying little more 
than half of postero-dorsal margin, anterior plate approximately half 
length of posterior, hinge teeth acute chevron-shape, relatively few 
in number, up to 18 in posterior series and 16 in anterior; ligament 
internal, oblique, posterior to umbo; internal shell ridge extends 
from mid posterior margin to approximately opposite the mid-point 
of the posterior hinge plate between and parallel to the lines of the 



post-umbonal carinae.The maximum length of the present speci- 
mens is 15.3mm. 

In most specimens the posterior shell margin is damaged, often 
being markedly shortened and specimens frequently show regenera- 
tion of the shell posterior to the posterior adductor. The repaired 
shell is without concentric ornamentation. It is possible that ex- 
tended siphons are predated upon and that the shell is damaged when 
this occurs. 








Fig. 40. Propeleda carpenteri. External lateral view of the right side of a shell and an internal lateral view of a right valve from Atlantis II station 237 
from the Argentine Basin. Scale = 1mm. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



127 




Fig. 41. Propeleda carpenteri. External lateral views of the right sides of 
two small shells to show differences in shape from the specimen 
illustrated in text-figure 40. Specimens taken from Atlantis II station 239 
from the Argentine Basin. Scale = I mm. 

The concentric ridges on the shell of the present specimens while 
not particularly marked are more so than those described by Dall, 
though the syntypes that we have examined are dead valves that are 
somewhat worn (Fig. 39). Our specimens also are very slightly more 
anteriorly extended than the syntypes, though the characteristic 
antero-dorsal curvature of the shell margin is the same. Our speci- 
mens correspond well with the figures given by James ( 1 972, figs 67 
and 68). These latter come are from similar depths (2340-2627m) to 
some of our own. It would appear that specimens from mid to lower 
slope differ slightly from those taken at shallower depths however, 
we regard the differences as being at most infrasubspecific. 

There is a marked change in shape during growth. Juveniles are 
much shorter than the adults and subsequent growth involves in- 
creasing elongation of the post-umbonal shell. The prodissoconch is 
extremely large measuring 630um in length. 



Internal morphology (Fig.42) 

The adductor muscles are oval in cross-section, the posterior being 
the more fusiform, both are set some distance in from the shell 
margin. The posterior adductor muscle lies opposite the limit of the 

posterior hinge plate, the anterior muscle is attached to the shell just 
dorsal of the mid-horizontal shell axis. The siphonal embayment is 
elongate, the siphons are slender and entire. The anterior sense organ 
is small in size. 

The foot lies in the anterior half of the mantle cavity in preserved 
specimens, it is relatively elongate and has numerous small papillae 
present along the margins of the sole. The palps are small, each 
bearing an extremely long, narrow, palp proboscis. Depending on 
the size of the specimen there are up to 17 palp ridges. The gills are 
elongate, slender, and have up to 17 plates. 

The course of the hind gut is similar to that in Nuculana in that it 
passes to the right side of the body where it forms a broad loop that 
passes close to the posterior wall of the anterior adductor muscle. 
The stomach is large and occupies a vertical position in the posterior 
part of the visceral mass. The digestive gland is extensive occupying 
much of the antero-dorsal visceral space. 

Propeleda louiseae (Clarke 1961) 

Type specimen. Hoiotype, MCZ 224958. 

Type locality. R. V. Vema biology station 121, Argentine Basin, 
1000 miles ESE of Buenos Aries, 5105 metres. 

Cited specimens. BMNH 1995058 

Nuculana (Thestyleda) louiseae Clarke 1961, 375, pl.l, fig. 7. 

Material: 



Cruise 



Sta Depth No Lat 
(m) 



Long 



Date Gear 



ARGENTINE BASIN 
Atlantis II 242 4382- 25 
60 4402 

243 3815- 2 

3822 
247 5208- 2 

5223 
256 3906- 3 
3917 



38"16.9'S 51"56.1'W 13.3.71 ES 

3736.8'S 52"23.6'W 14.3.71 ES 

43"33.0'S 48*58. l'W 17.3.71 ES 

37"40.9'S 52*19.3'W 24.3.71 ES 




Fig. 42. Propeleda carpenteri. Internal morphology as seen from the right side of a specimen taken from Atlantis II station 239 from the Argentine Basin. 
For identification of the parts see text-figure 7, p. 106. Scale = 0.5mm. 



128 



J.A. ALLEN AND H.L. SANDERS 




Fig. 43. Propeleda louiseae. a & b, external lateral views of the right sides of shells of differing size to show differences in shell proportions with growth; 
note outline internal morphology through semi-transparent shells, in particular the form of the hind gut and position of the adductor muscles; c, outline of 
shell from the left side showing the outline of hind gut and adductor muscles; d, internal view of a left valve. All specimens taken from Atlantis II station 
242 from the Argentine Basin. Scales = 1mm. 



Type specimen examined by HLS. 

Distributed at abyssal depths in the Argentine Basin; depth range: 
3815-5223 metres. 

Shell description (Fig.43) 

Shell extremely elongate, slender, fragile, semi-transparent, 
ornamented with moderately spaced concentric ridges, two post- 
umbonal rounded ridges, one dorsal and one ventral at posterior 
shell margin and crossed vertically by wavy continuations of the 
concentric ridges, faint anterior radial ridge from umbo to antero- 
ventral margin; umbo slightly raised, far anterior (post-umbonal 
length 65-70% of total length), inwardly facing; antero-dorsal 
margin with short proximal notch, distally margin raised and slightly 
concave, faint angulation before anterior margin, anterior margin 
joins with ventral margin in smooth curve, postero-ventral margin 
very slightly sinuate, posterior margin angled and sinuate, postero- 
dorsal margin notched at umbo, distally somewhat raised and keeled, 
concave overall but slightly sinuous dorsal to hinge plate; hinge 
plate relatively broad, posterior plate short occupying approxi- 
mately half the postero-dorsal shell margin, hinge teeth elongate, 
acute chevron shape, up to 12 in anterior and 20 in posterior series; 
ligament small, internal, oblique, pear-shaped; rounded internal 
ridge extends from umbo to posterior margin and marks junction 
between inhalent and exhalent siphons. The maximum length of the 
present specimens is 20.3mm. The prodissoconch is large measuring 
300um in length. 

Clarke (1961) records a long, thin, external ligament in the type 



specimen - the latter being a single large valve. We find no evidence 
of an external part to the ligament and believe that Clarke mistakenly 
confused periostracum along the postero-dorsal margin for liga- 
ment. 

Internal morphology 

With one notable exception the internal morphology of Plouiseae 
differs little from that of P. carpenteri. 

Unlike the latter species the hind gut of P. louiseae first passes to 
the left side of the body where it forms a relatively small loop 
immediately ventral to the umbo (Fig.43). From there it passes to the 
right of the body and forms a loop that is considerably larger than 
that on the left although not as extensive as that in P. carperteri (Figs 
42&43). 

The adductor muscles are relatively large, the elongate poste- 
rior muscle is situated at the distal limit of the posterior hinge 
plate. The gill is very short and slender with few (c. 13) gill- 
plates. 

Propeleda paucistriata (new species) 

Type SPECIMEN. Holotype BMNH 1995059; Paratypes BMNH 
1995060. 

TYPE LOCALITY. Atlantis II station 203, Angola Basin, 08°48.00'S 
12°52.00'E, 527-542m. 






DEEP-SEA PROTOBRANCHIA (BIVALVIA) 

Material: 



Cruise Sta 


Depth No 

(m) 


Lat 


Long 


Date 


Gear 


ANGOLA BASIN 
Atlantis II 203 

42 


527- 31 
542 


08"48.00'S 


12"52.00E 


23.5.68 


ES 



Distribution. Restricted to the Angola Basin at upper slope 
depths, 527-542 metres. 

Shell description (Fig.44) 

Elongate, fragile, transluscent shell, moderately slender, two cari- 
nate ribs from umbo to posterior margin, widely spaced prominant, 
relatively broad, concentric ribs with overhanging ventral margin, 
2—4 fine concentric lines between ribs, between carinae vertical ribs 
and lines equally prominant; umbo moderately raised, far anterior in 
largest specimens (post-umbonal length 79% of total length) but less 
so in smaller specimens, beaks inwardly facing; antero-dorsal mar- 
gin sloping, proximally convex but almost straight in small specimens, 
joins with anterior and antero-ventral margins in smooth curve; 
postero-ventral margin very slightly sinuous, posterior margin usu- 
ally damaged in large specimens, intact margin angled and sinuate, 
forming a hook dorsally where postero-dorsal margin and dorsal 
carina meet, postero-dorsal margin concave, proximally raised, 
elongate escutcheon outlined by dorsal carina; hinge elongate, 
moderately broad, large, acute chevron-shaped teeth up to 16 in 
anterior series and up to 28 in posterior series, anterior series extends 
to anterior limit of anterior adductor muscle, posterior series extends 
approximately half length of postero-dorsal margin to anterior limit 



129 

of posterior adductor muscle, ventral margin of hinge plate corre- 
sponds to line of dorsal carina, ventral to umbo teeth approach shell 
margin, anterior and posterior hinge plates continuous; ligament 
internal, ventral to umbo and close to shell margin, slightly inclined 
posteriorly; rounded internal ridge extends from umbo to posterior 
margin. The maximum recorded shell length is 14.1mm. The 
prodissoconch is very large and measuring 560pm in length. 

Juvenile shells are more ovate and, before posterior elongation 
occurs, could be mistaken for a yoldiellid (Fig.44). 

Internal morphology (Fig.45) 

The anterior adductor muscle is oval in cross section, while the 
posterior adductor is smaller and more elongate. Both are set in from 
the shell margin, the posterior is positioned at approximately two- 
thirds the distance between the umbo and the posterior limit of the 
shell. There is a small anterior sense organ formed from the sensory 
fold of the mantle, ventral to the anterior adductor. The siphons are 
joined with their ventral margins fused to form entire lumina. They 
are slender and particularly elongate and when contracted are con- 
tained in the elongate siphonal embayment. The foot and viscera lie 
in the anterior half of the mantle cavity. The foot is elongate and 
directed anteriorly. In most preserved specimens the tip of the foot 
lies between the anterior adductor and the shell margin. The margins 
of the sole are fringed with numerous relatively small papillae. 
There are three anterior and two posterior pedal retractor muscles. 
The palps and gills are markedly narrow and elongate. The are at 
least 22 palp ridges in the largest specimens and the palp probosci- 
des are attenuate each with a straight dorsal margin and a papillate 
ventral margin. In a few preserved specimens the palp proboscides 
extend from the feeding aperture. The gills are similarly attenuate 




Fig. 44. Propeleda paucistriata. External lateral views of the right sides of four shells of differing size to show change of shape with growth and an 
internal view of a right valve. All specimens taken from Atlantis II station 203 from the Angola Basin. Scale = 1mm. 



130 



J.A. ALLEN AND H.L. SANDERS 






Fig. 45. Propeleda paucistriata. a, internal morphology as seen from the right side; b, part of the left side of the same specimen to show details of the 
course of the gut; c, the internal morphology as seen from the left side of a much larger specimen. All specimens taken from Atlantis II station 203 from 
the Angola Basin. For identification of the parts see text-figure 7, p. 106. Scales = 1mm. 



and extend from the posterior visceral mass to the anterior limit of 
the posterior adductor. There are at least 22 gill plates in larger 
specimens. A slender extension of the axis extends from each gill 
from ventral to the posterior adductor to the inner junction between 
inhalent and exhalent siphons. 

From the large combined stomach and style sac the course of the 
hind gut takes it first dorsal and posterior to the stomach and then to 
the left side of the body where it makes a small loop. From there it 
passes ventral to the umbo to the right side of the body where it 
makes a much larger loop at the perimeter of the viscera and passing 
close to the inner face of the anterior adductor. From there it passes 
mid-dorsally to the anus. The mouth is set some distance posterior to 
the anterior adductor muscle. The oesophagus is broad and elongate 
and the combined stomach and style sac is positioned vertically in 
the posterior part of the visceral mass. The pedal ganglion is large 
and lies immediately anterior to the junction of mid gut and hind gut. 

The shell surface in some larger specimens is covered with 
epifaunal solitary hydroids. This would indicate that P. paucistriata 
lives close to the surface of the sediment. This is also suggested by 
the fact that the posterior tips of the shells of larger specimens are 
broken. We believe that this is the result of predation on the siphons. 

The extreme post-adductor elongation of the shell is advanta- 
geous in that it provides distance between predator and the more 
vulnerable viscera with damage being restricted to more easily 
generated tissue. 

The shell of this species differs from others described by the small 
number of pronounced concentric shell ridges and we name it with 
reference to this characteristic feature. This is the first species of 
Propeleda to be recorded off the south-west coast of Africa. At 
approximately the same latitude off the east coast of Natal a species 
named Leda lanceta by Boshoff ( 1 968) occurs at upper slope depths. 
Nijssen-Meyer ( 1 972) believes that this latter species is aPropeleda, 



and we concur with her. P. lanceta is more robust, more arcuate and 
with far more numerous concentric ridges than is the case in P. 
paucistriata. 



DISCUSSION 

The major point of interest in this particular account of deep-sea 
protobranch bivalves is the evidence it provides to further our 
understanding of the evolution of the nuculanoid protobranchs. In 
our earlier studies on the Tindariidae we speculated as to how the 
nuculanoids could have evolved from the nuculoids (Sanders and 
Allen, 1977). In functional terms, this involved a change in the 
inflow of water into the mantle cavity from an antero-ventral posi- 
tion to a posterior position and the begining of specialization of the 
posterior mantle edge, a view also expressed by Yonge (1939). In the 
tindariids this latter involves the development of sensory papillae 
from the sensory fold of the mantle at the points of ingress and exit 
of the circulatory water. Although infaunal and deposit feeding, the 
tindariids, like the nuculids, live close to the surface and, like many 
other bivalves that occupy this position, they are ovate and robust. 
Many of these subsurface dwelling bivalves, including the tindariids, 
have stout external ligaments. 

The development of siphons was the next step in the evolutionary 
process and the neilonellids are illustrative of this. The shell form and 
ligament as seen in the tindariids is largely retained in the neilonellids, 
but short siphons, as yet only fused dorsally, are now present and these 
are contained in a shallow siphonal embayment. Although the shell 
remains stout and ovate there is some posterior elongation and an area 
ventral to the inhalent siphon from where the palp proboscides are 
extended is more defined. Like the tindariids the neilonellids are 
deposit feeders living close to the surface of the sediment. 



DEEP-SEA PROTOBRANCHIA (BIVALVIA) 



131 



The hind gut mNeilonella, like that in Tinclaria, has a wide lumen 
and single pronounced typhlosole. Although the course that the hind 
gut takes in neilonellids makes a single loop on the right side of the 
body, it does not penetrate mantle space as it does in tindariids 
(Sanders and Allen, 1977). In this respect the hind gut of neilonellids 
probably represents the more primitive condition. We have argued 
elsewhere (Allen, 1992) that elongation and the complexity of form 
of hind gut configuration are related to food procurement at great 
depths, and this applies to the tindariids (Sanders and Allen, 1977). 
The neilonellids are for the most part upper slope species and the 
hind gut would be expected to be less specialized and less elongate. 

In Nuculana posterior elongation becomes more extreme and the 
ventral margins of the combined siphons are fused such that the 
exhalent and inhalent lumena are separate and entire. The shell 
remains robust, but is more slender The ligament is restricted to a 
small internal structure separating elongated anterior and posterior 
series of hinge teeth. We believe that elongation is correlated to the 
almost vertical orientation of the animal in the sediment but which 
retains contact with the surface via the extended posterior body and 
siphons. The genus Nuculana is found mainly in shelf and upper 
slope sediments and as such the available food resources are rela- 
tively abundant. The hind gut is not greatly extended and remains as 
a single loop to the right side of the body. 

In Propeleda the evolutionary trend of posterior body elongation 
seen in Nuculana becomes is more extreme, particularly posterior to 
the posterior adductor muscle. The posterior adductor muscle is 
more elongate and dorso-ventrally narrow, and the gill, gill axes, 
siphons and the palp proboscides are exceptionally long and slender. 
The shell of Propeleda, particularly in abyssal species, is much more 
fragile and is further specialized in that it possesses an internal 
posterior longitudinal ridge. The function of this ridge is not entirely 
clear and has await examination of the living animal but, possibly, it 
is involved in the separation of excretion, feeding and respiratory 
functions in the extremely elongate posterior mantle cavity. It may 
also help to strengthen the otherwise very fragile shell and assist in 
predation damage limition. In Propeleda post-adductor elongation 
involves body tissues that can be relatively easily regenerated, much 
in the same way as has been reported in deposit feeding tellinids 
(Edwards, Steele and Trevallion, 1970). Specimens showing shell 
repair posterior to the posterior adductor are present in our samples. 

The evolution of the Ledellinae and an assessment of their func- 
tional morphology was discussed earlier (Allen and Hannah, 1989). 
In respect of the species of Ledella and Tindariopsis described here, 
little needs to be added to that account other than to note, again, that 
the hind gut in these abyssal protobranchs is extraordinarily length- 
ened and takes the most complex courses within the visceral mass. 

The other item of note is the description of yet another ledellid in 
which the shell, after reaching a certain length, changes its direction 
of growth. In Ledella aberrata as in L. ultima the result of this 
change is to produce a broad shell margin and lateral expansion of 
the shell cavity. This adaptation has been construed as possibly 
providing more space for the gonads that begin to develop at about 
the time the change in direction occurs. 

Acknowledgements. It is fitting that in this paper we thank our numer- 
ous friends and colleagues, particularly French colleagues of the Biogas and 
Walda cruises, who provided much material and gave much advice and 
encouragement. We would like to mention three colleagues by name. George 
Hampson of the Woods Hole Oceanographic Institution and Fiona Hannah 
(Lonsdale) of the University Marine Biological Station, Millport, who have 
given us tremendous close support over the years and without their help it is 
likely that we would not have succeeded in this task within our lifetimes, and 
Kathy Way of the Natural History Museum, London, who has been so quick 



in responding to our requests for help particularly in our quest for early 
literature and specimens for comparison. Much of the work was supported by 
grants from the Natural Environment Research Council. 



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Title. The title page should be arranged with the full title; name(s) 
of author(s) without academic titles; institutional address(es); sug- 
gested running title; address for correspondence. 

Synopsis. Each paper should have an abstract not exceeding 200 
words. This should summarise the main results and conclusions of 



the study, together with such other information to make it suitable 
for publication in abstracting journals without change. References 
must not be included in the abstract. 

Text. All papers should have an Introduction, Acknowledgements 
(where applicable) and References; Materials and Methods should 
be included unless inappropriate. Other major headings are left to 
the author's discretion and the requirements of the paper, subject to 
the Editors' approval. Three levels of text headings and sub-headings 
should be followed. All should be ranged left and be in upper and 
lower case. Supra-generic systematic headings only should be in 
capitals; generic and specific names are to be in italics, underlined. 
Authorities for species names should be cited only in the first 
instance. Footnotes should be avoided if at all possible. 

References. References should be listed alphabetically. Authori- 
ties for species names should not be included under References, 
unless clarification is relevant. The author's name, in bold and lower 
case except for the initial letter, should immediately be followed by 
the date after a single space. Where an author is listed more than 
once, the second and subsequent entries should be denoted by a long 
dash. These entries should be in date order. Joint authorship papers 
follow the entries for the first author and an '&' should be used 
instead of 'and' to connect joint authors. Journal titles should be 
entered in full. Examples: (i) Journals: England, K.W. 1987. Certain 
Actinaria (Cnidaria, Anthozoa) from the Red Sea and tropical Indo- 
Pacific Ocean. Bulletin of the British Museum (Natural History), 
Zoology 53: 206-292. (ii) Books: Jeon, K.W. 1973. The Biology of 
Amoeba. 628 p. Academic Press, New York & London, (iii) Articles 
from books: Hartman, W.D. 1 98 1 . Form and distribution of silica in 
sponges, pp. 453^493. In: Simpson, T.L. & Volcani, B.E. (eds) 
Silicon and Siliceous Structures in Biological Systems. Springer- 
Verlag, New York. 

Tables. Each table should be typed on a separate sheet designed to 
extend across a single or double column width of a Journal page. It 
should have a brief specific title, be self-explanatory and be supple- 
mentary to the text. Limited space in the Journal means that only 
modest listing of primary data may be accepted. Lengthy material, 
such as non-essential locality lists, tables of measurements or 
details of mathematical derivations should be deposited in the 
Biological Data Collection of the Department of Library Services, 
The Natural History Museum, and reference should be made to 
them in the text. 

Illustrations. Figures should be designed to go across single 
(84 mm wide) or double (174 mm wide) column width of the 
Journal page, type area 235 x 174 mm. Drawings should be in black 
on white stiff card or tracing film with a line weight and lettering 
suitable for the same reduction throughout, either 50%, 30% or 
25%. After reduction the smallest lettering should be not less than 
10 pt (3 mm). All photographs should be prepared to the final size of 
reproduction, mounted upon stiff card and labelled with press-on 
lettering. Components of figure-plates should be abutted. All fig- 
ures should be numbered consecutively as a single series. Legends, 
brief and precise, must indicate scale and explain symbols and 
letters. 

Reprints. 25 reprints will be provided free of charge per paper. 
Orders for additional reprints can be submitted to the publisher on 
the form provided with the proofs. Later orders cannot be accepted. 



71 Indian Ocean echinoderms collected during the Sindbad Voyage (1980-81): 3. Ophiuroidea 

and Echinoidea 

Andrew R.G. Price and Francis W.E. Rowe 
83 Rare cyclopoid copepods (Crustacea) from Mediterranean littoral caves 

D. Jaume and G.A. Boxshall 
101 Studies on the deep-sea Protobranchia (Bivalvia): the family Neilonellidae and the family 

Nuculanidae. 

J. A. Allen and H.L Sanders 



ZOOLOGY SERIES 

Vol. 62, No. 2, November 1996